Mendel University in Brno

Faculty of Forestry and Wood Technology

Plant Botany
An introduction to plant anatomy, morphology and
physiology

Milena Martinková, Martin Čermák, Roman Gebauer, Zuzana Špinlerová

Brno 2014
Tato skripta byla vytvořena v rámci projektu InoBio – Inovace biologických a lesnických disciplín pro
vyšší konkurence schopnost, registrační číslo projektu CZ.1.07/2.2.00/28.0018. za přispění finančních
prostředků EU a státního rozpočtu České republiky.

Translated from the Czech edition "Kolařík et al. (2010) Péče o dřeviny rostoucí mimo
les - II." published by ČSOP Vlašim.

Translated by Ing. Roman Gebauer Ph.D. and Mgr. Lucie Novotná

Authors ´contribution:

Milena Martinková (Chapter 1, 2, 3, 7 and 8)

Roman Gebauer (Chapter 4)
Martin Čermák (Chapter 5)
Zuzana Špinlerová (Chapter 6)

Author of figures - Milena Martinková (author of figures 9, 14, 15, 16, 18, 20 and 21 is
Roman Gebauer)
Contents

1 Introduction ...............................................................................................................3
1.1 Life domains and phylogeny of tree growth on Earth ........................................ 3
1.1.1 Prokaryote .................................................................................................. 3
1.1.2 Eukaryote ................................................................................................... 4
1.1.3 Evolutionary history of vascular plants ....................................................... 4
1.1.4 General characteristics and types of trees ................................................. 6
1.1.5 What are the benefits of trees?................................................................... 8
1.1.7 Tree reaction on the unfavourable condition .............................................. 9
1.1.8 Ontogeny of tree growth ........................................................................... 10
1.1.9 Dynamic of tree activity ............................................................................ 12
2 Plant Cell ................................................................................................................13
2.1 Membrane complex, semi-autonomy organelles and cytoskeleton ................. 14
2.2 Nucleus ........................................................................................................... 15
2.3 Cell ontogenesis.............................................................................................. 16
3 Tissues ...................................................................................................................18
3.1 The origin of primary meristem........................................................................ 20
3.2 Primary meristems .......................................................................................... 21
3.3. Permanent primary tissues ............................................................................ 23
3.3.1 A system of primary surface tissues ......................................................... 23
3.3.2 A system of primary basic tissues ............................................................ 24
3.3.3 A system of vascular tissues ................................................................... 25
3.4 Secondary meristems ..................................................................................... 25
3.4.1 Cambium .................................................................................................. 25
3.4.2 Phellogen.................................................................................................. 28
4 Roots ......................................................................................................................30
4.1 The origin of root ............................................................................................. 30
4.1.1 General characteristics of root systems in trees ....................................... 31
4.2 Root functions ................................................................................................. 33
4.3 Root zone activity (primary growth) ................................................................. 33
4.3.1 Meristem zone of the root primary growth ................................................ 33
4.3.2 The elongation zone ................................................................................. 34
4.3.3 Differentiation zone................................................................................... 34
4.4 Secondary growth and root functions .............................................................. 36
4.5 Mycorrhiza ...................................................................................................... 37
4.5.1 Vesicular-arbuscular mycorrhiza (VAM) = arbuscular mycorrhiza (AM) ... 38
4.5.2 Ectomycorrhiza (EM) ................................................................................ 39
 4.5.3 Ectendomycorrhiza ................................................................................... 39
4.6 Metamorphoses of root ................................................................................... 40
4.7 Limiting factors of root system growth and functions....................................... 40
5 The stem.................................................................................................................42
5.1 Secondary growth of stem .............................................................................. 43
5.2 Development of annual rings .......................................................................... 44
5.3 Wood of coniferous species ............................................................................ 45
5.4 Wood of deciduous species ............................................................................ 46
5.5 Stem morphology ............................................................................................ 47
5.6 Bud.................................................................................................................. 49
6 Leaf.........................................................................................................................51
6.1 Phyllogenesis – the evolution of a leaf ............................................................ 51
6.2 Ontogenesis – the individual development of a leaf ........................................ 51
6.3 The arrangement of leaves on a stem............................................................. 53
6.4 The outer structure of assimilation leaves ....................................................... 54
6.5 The inner leaf structure ................................................................................... 56
6.6 The inner needle structure .............................................................................. 60
6.7 Evaluation of foliage state ............................................................................... 61
6.8 Leaf metamorphoses ...................................................................................... 61
7 Propagation and reproduction of woody plants .......................................................61
7.1 Means of reproduction and types of diasporas................................................ 62
7.1.1 Artificial vegetative reproduction ............................................................... 62
7.2 Plant life cycle and the significance of seeds in phylogenesis of plants .......... 63
7.2.1 Dissemination by diasporas of a generative type ..................................... 64
8 Basics of woody plant physiology ...........................................................................71
8.1 Photosynthesis ................................................................................................ 72
8.1.1 The impact of exogenous and endogenous factors on photosynthesis .... 75
8.1.2 CO2 exchange in plants ........................................................................... 76
8.1.3 Photosynthesis and respiration................................................................. 77
8.2 Respiration ...................................................................................................... 77
8.2.1 The process of respiration - oxidation of glucose ..................................... 79
8.2.2 Factors of respiration ................................................................................ 81
8.3 Photorespiration .............................................................................................. 82
8.4 Water regime of woody plants......................................................................... 83
8.4.1 Water content and its thermodynamic state.............................................. 84
8.4.2 The movement of water in a cell ............................................................... 86
8.4.3 Water saturation deficit ............................................................................. 87
8.4.4 Water potential of plants ........................................................................... 87
 8.4.6 The uptake and conduct of water ............................................................. 87
8.4.7 Release of water by plants ....................................................................... 91
8.4.8 Water balance in a plant ........................................................................... 96
8.5 Mineral nutrition of woody plants and the significance of nutrients.................. 96
8.5.1 Roles of individual nutrients ...................................................................... 98
8.5.2 Ion relations .............................................................................................. 99
8.5.3 Nutrition disorders ...................................................................................100
1 Introduction

If we wish to grow a plant we should perceive it in relation to its position in the

vegetation system—it informs us on the stage of development in the long history of
life on Earth. Moreover, we should be aware of its demands on the environment
(i.e. environmental requirements), the degree of adaptability to changes in the
environment, sensitivity or resistance to insect, fungal and other pathogens. We
should be informed about the natural extension of this taxon on Earth (whether it is
a native species or naturalized with us, or a newly introduced species, with which we
lack experience) and also the origin of its direct ancestors (whether they are
individuals originating from seed, clones or grafts). Simply put, you need to
understand each plant grown in time and space and, what is more, as these are
living organisms, we should take into consideration their life experience and
reactions—absolute and relative age (i.e. ontogeny stage) and their growth rates
and changes in shape.

1.1 Life domains and phylogeny of tree growth on Earth

Life on Earth is composed of unicellular and multi-cellular living systems

(organisms capable of performing all life functions and information flow). Cellular
systems originate in one hypothetically assumed ancestor called progenot, which
evolved in two evolutionary lines about 3.5 billion years ago. One directed to the
bacteria (Prokaryote) domain and the second to the Eukaryote domain.

1.1.1 Prokaryote

Bacteria and archaea domains are unicellular organisms of prokaryotic type.

Their cells contain cytoplasm, plasma membrane and nucleoid. Nucleoid is not
divided from the cytoplasm by a membrane; it is composed of one, generally a
circular, double-stranded piece of deoxyribonucleic acid (DNA). The interior of
these cells is not divided by a system of membranes (mitochondria and plastids
are missing). Prokaryotic organisms are capable of heterotrophic nutrition (as
saprophytes and parasites growing both in the aerobic or anaerobic environment) as
well as autotrophic nutrition (chemo-or phototrophic). They include species
producing methane, species tolerant of high temperatures (up to 113 °C), high
pressure, salt and acid environment. They have a relatively large surface contact
with the environment, a rapid exchange of matter and energy and the fastest
metabolism (generation time, i.e. the time at which parental generation gives rise a
new generation, may only take 30 minutes, they are able to synthesize up to
15 amino acids in one second). They make up about one half of the biomass
(living matter) of the world.
There are typically 40 million bacteria cells in a gram of soil and a million
bacteria cells in a millilitre of fresh water. They live in water, in bodies of other
organisms, and can spread through the air. They are mainly found in the soil, where
they take part in mineralization of remains of plant and animal bodies. Through
the decomposition of their proteins ammonia is released, and ammonium salts are
then oxidized by nitrifying bacteria into nitrites and nitrates, and these are the source
of plant nutrition. Some nitrogen-fixing bacteria such as Azotobacter and Rhizobium
are involved in the nitrogen cycle by binding to their proteins and nucleic acids of
atmospheric nitrogen. Azotobacter lives and fixes nitrogen independently of other

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organisms. On the other hand, Rhizobium fixes nitrogen after entering root nodules of
legume plants. Conversely, denitrifying bacteria deprive soil of nitrates and nitrites as
they reduce them to molecular N2.
Bacteria are also involved in the carbon cycle, the source of carbon dioxide
(CO2). CO2 is released into the atmosphere by the oxidization activity of organisms
(respiration of plants, animals, fermentation of chemoheterotrophous bacteria). The
released CO2 is a source of carbon for all photoautotrophic organisms, especially in
the subkingdom Viridiplantae – green plants. Another element whose cycle is
affected by the bacteria is sulphur. Sulphur is contained in many proteins in plant and
animal bodies. Saprophytic bacteria release sulphide from their residues utilized by
phototrophic sulphur bacteria as a source of hydrogen. Sulphur is then oxidized to
sulphate by chemoautotrophous bacteria, which is available to plants. Green plants
are of a fundamental importance to heterotrophic consumers as essential
primary producers of organic matter.

1.1.2 Eukaryote

The most evolutionarily advanced organisms belong to eukaryotic domain,

whose minimum building unit, capable of all basic life functions, is a eukaryotic
cell – i.e. cell where the information molecule DNA is localized in the nucleus
enclosed by a membrane. The most important evolutionary branches include
eukaryotic protozoa and chromista such as soil organisms, fungi, animals as
heterotrophic organisms and plants, mainly as producers of organic matter, i.e.
autotrophic organisms.

1.1.3 Evolutionary history of vascular plants

Once plants had reached the land, there were two approaches to dealing with
desiccation. The bryophytes avoid it or give in to it, restricting their ranges to moist
settings, or drying out and putting their metabolism "on hold" until more water arrives.
Tracheophytes resist desiccation: they all bear a waterproof outer cuticle layer
wherever they are exposed to air (as some bryophytes do), to reduce water loss,
but—since a total covering would cut them off from CO2 in the atmosphere—they
rapidly evolved stomata, small openings allowing for the gas exchange.
Tracheophytes also developed vascular tissue to aid in the movement of water
within the organisms and moved away from a gametophyte dominated life cycle.
Vascular tissue also facilitated upright growth without the support of water and paved
the way for the evolution of larger plants on land.

1. Cooksonia, Rhynia and Psilophyton are the first known vascular plants
(Tracheophytes). They lived from the upper Silurian to early Devonian (443–
393 million years ago) and they were leafless and rootless plants consisted of stem
tissue which branched along an axis. Their bodies already had an epidermis with
stomata and these vascular plants had ingeniously invented a chemical called lignin
that toughens the walls of plant cells. The earliest documented plant of this type is
Cooksonia, but a complete fossil material was not found yet. On the other hand,
complete fossils were found for Rhynia plants. The axes of Rhynia exhibit a
maximum diameter of 3 mm and the plant probably attained a height of up to 20 cm.
Psilophyton is a genus with more complex structure than any other plants of
comparable age (e.g. Rhynia) and is thought to be part of the group from which the

4
modern ferns and seed plants evolved. P. forbesii is the largest documented species
of Psilophyton. Reconstructions suggest that they were around 60 cm tall and
equipped with more advanced type of tracheid. However, no species of this family
reached tree growth because of isometric growth (i.e. almost uniform in all
directions), which would not be functionally and mechanically sustainable in larger
sizes.
2. Pteridophytes (clubmosses, horsetails, ferns and progymnosperms) is group of
vascular plants where diploid sporophyte (with a double set of chromosomes in the
cell nuclei) predominates over gametophyte, which is haploid (with one set of
chromosomes) in individual lives. Trees (with a minimum height of 4 m and diameter
of 7.5 cm – measured at breast height, i.e. 1.3 m above ground) appeared for the first
time in the history of Earth flora between the middle and late Devonian (350 million
years ago). Since then they have been gaining in size in all dimensions, mainly in the
stem diameter. The earliest known trees originated in the Wattieza genus. At those
times, lush and dense vegetation spread in humid habitats, while dry habitats with a
low ground water only had a sparse vegetation cover. The plants growing in wet
habitats were competing for light and this selective pressure was responsible for the
evolution (phylogeny) of tree habitus. The tree-high growth was discovered
independently, but more or less at the same time in ferns (Pseudosporochnus,
Calamophyton), clubmosses (Protolepidodendropsis, Cyclostigma), horsetails
(Psudobornia) and progymnosperms (Eospermatopteris, Archaeopteris). At the same
time tree plants, especially progymnosperms, (genus Duisbergia) occurred in drier,
sparsely populated habitats, too. In this case, the tree growth was advantageous
in terms of improving bioclimate in the crown (to avoid temperature fluctuations
near the soil surface) and in terms of ability to limit water loss (due to massive solid
trunk, which also served as a reservoir of water and other substances). Furthermore,
trees had an advantage of producing high-quality organic matter (mineralization of
dead tree parts brings nutrients back into the soil in the most favourable ratio and
form for a particular species), long life and slow population growth, which was an
asset to sustaining stability in an individual as well as the whole forest. Yet, trees
could appear in evolution only after satisfying their basic living need. It needed
sufficiently developed and deep soil in order to provide a tree with solid anchoring
and covering of their needs for water and mineral nutrients.
3. Gymnosperms plants (conifers, cycads, Ginkgo, seed fern and Gnetales) as a
group of seed-producing plants appeared in the Palaeozoic Devonian 405–
345 million years ago. They have an imperfect protection of ovules that grow
freely on scales. These ovules ripen into seeds. Pollen is transferred directly onto
the naked ovules. Male and female micro and megasporophylls usually form
separated male and female cones. The pollination by wind occurs. Seed ferns are an
extinct group of plants with tree or liana structures, large pinnate leaves resembling
fern leaves. The most primitive group of present seed plants are Cycads,
nowadays found in tropical and subtropical areas reaching about 130 species. A
class of Ginkgo originated in late Paleozoic and gradually spread to a large areal of
the Earth. In the Tertiary (65–2.5 million years ago), however, most Ginkgo species
died out, and only a single species – Ginkgo biloba survived till present flora. The first
conifers have been known since the Carboniferous period (345–280 million years
ago). Their greatest era was the Mesozoic (252–66 million year ago). At present
coniferous forests cover approximately one third of all forest ecosystems area.
4. Angiosperms (Flowering plants – Magnoliophyta – dicots and monocots)
appeared at the end of the Mesozoic in Late Cretaceous (140 million years ago).

5
Angiosperms differ from their predecessors by the position of ovules inside the
carpel (gynoecium), enclosed seeds in fruits and stamens with carpel that are
usually protected by colourful corolla, which attracts pollinators. Angiosperms
offsprings of the Cretaceous era are found nowadays in tropical and subtropical
areas (myrtle, Magnolia or Liriodendron). In higher latitudes, this group is represented
by mild climate species, such as willow, oak, birch, beech. The treelike monocot
species do not have a continuous and full wood stem cylinder; stem grows as a set
of leaf petioles and sheaths with a possible coappearance with adventitious roots.
Conductive systems are built from closed vascular bundles, which are composed of
xylem and phloem. The xylem typically lies adaxial with phloem positioned abaxial.

1.1.4 General characteristics and types of trees

The prerequisites for the existence of trees do not only include the capability to
build a functional multicellural organism (e.g. vascular architecture), but also to
synthesize the necessary construction, storage, defence and regulatory
substances. These include accumulation of the storage and other substances into
the cells (starch, proteins, fats or substances of the defence system – anthocyanins,
tannins, dyes, aromatic and bitter substances, saponins, glycosides, alkaloids or
metabolic waste products – crystals of calcium oxalate and calcium carbonate etc.),
into the cell walls (cellulose, hemicellulose, pectin substances, cutin and other high
molecular weight waxes, lignins, suberin, polysaccharide mucilaginous substances
etc.) or into specialized locations (resins, essential oils, latex, gum, rubber and many
others).
Woody plants are perennial plants whose woody stem lives for at least two
growing seasons (e.g. raspberry, blackberry), but usually longer, i.e. from many
years, exceptionally up to thousands of years. Woody plants are studied by
dendrology (from Greek: dendron – meaning a tree; logia – science or study).
However, dendrology does not cover the whole part of the system (unlike the case of
mycology – that studies exclusively fungi), since a lot of systematic units include both
herbs and woody plants (e.g. within the Ranunculaceae family fig buttercup Ficaria
verna is a typical herb and old man’s beard Clematis vitalba is a woody lian or within
the genus of Sambucus there is a herbaceous species dwarf elder Sambucus ebulus,
while individuals of elderberry Sambucus nigra grow as woody plants – shrub or tree.
Moreover, there are a number of herbs able to build woody stems strong and solid
enough and lateral branches that structurally correspond to woody species
(wormwood, Atriplex, hops) – but these species are not considered as trees, since
their above-ground systems die in the same year as they originate.
Woody plants – Plantae lignosae – are perennial plants with woody above-ground
and root systems, exclusively adapted to the terrestrial environment. They can be
divided into Holoxyles, whose stems lignify in the same growing season as they
elongate, and Hemixyles, whose stems fully lignify only as late as the following
season (i.e. in autumn their apical parts are still herbal pendant as they lack strength;
such as blueberry). If an unlignified part is damaged in rest season, this does not
present a grave loss to a plant. The opposite is true – if it survives, a hemixyle has an
advance when the photosynthesis starts at the beginning of the next growing season.
According to a plant design (physiognomy), woody plants can be distinguished as
trees (a distinct main trunk remaining unbranched in its lower part (the height may
vary depending on the species and conditions from just a few cm such as seedlings

6
or bonsai to more than 100 m), shrubs (higher than 0.8 m not developing a distinct
main trunk, with the stem branching of its basal part above or below the soil
surface).
Trees have a large, extensive root system, which passes into a compact, space-
minimized (intense) and clearly established trunk near the soil surface. A trunk
branches at a certain height above the ground and carries a crown. The crown
usually grows in its periphery, i.e. in height and width and the stem can be found
either around full length (e.g. spruce), or it may soon form strong, numerous skeletal
branches (as solitary broadleaf).
Shrubs, woody cushion plants and creeping woody plants also have extensive
above- and below-ground systems, but do not build a trunk—they develop new and
stronger branches from the lower buds, i.e. close to the soil surface. Shrubs with a
root system covered in the soil are known as chtonophytes. Above-ground parts of
woody cushion plants also have a nearly hemispherical shape with dense, short
shoots around the surface. The surface of such a compact unit is relatively small
related to the volume of the occupied space—such a shrub is thus protected from
temperature fluctuations and drying out. Above-ground parts of creeping woody
plants are pressed against the surface of the soil (e.g. cotoneaster) and the ends of
branches may be ascending.
Epiphytic shrubs grow on other plants, usually trees (epiphytes can also be currant,
yew tree and a number of other species and such trees as rowan or maple growing
on other trees and generally not forming a trunk in these unfavourable conditions).
They do not harm the host plant if there is enough space in the crown or if the host
plant falls and the epiphyte plant reaches the soil surface. Though, typical tropical
and subtropical epiphytes may outgrow a host tree and kill it (strangler).
Hemiparasitic shrubs such as yellow mistletoe and European mistletoe are
connected with their host plants directly by haustoria; they prosper in the top, sunlit
part of a tree crown, where their green leaves produce organic matter to their own
use (Fig. 1). They withdraw solutions from the host species that were obtained
with a great energy lost from the soil and pulled up against the force of gravity. These
shrubs often have a spherical shape whose surface is ideal for capturing solar
radiation and reducing excessive water loss by evaporation into the air.

Fig. 1 Oak branch infested by yellow mistletoe (Loranthus europeaus). The branch
top part dyed as nutrients and water were taken by mistletoe.

Woody lianas can be divided into several types:

7
• entwine – their stems perform rotational movements at their apices, which serve to
wrap around a support. Later they grow, get thicker and can graft among coils – e.g.
Dutchman’s pipe (Aristolochia macrophylla), Chinese wisteria (Wisteria sinensis)
• prop – often shortened reverse stems or emergencies – prickly outgrowths of bark,
which hold lians on the substrate. E.g. blackberry (Rubus fruticosus)
• with tendrils – such as grapevine (Vitis vinifera)
• with tendrils with suction pads, as Japanese ivy (Parthenocissus tricuspidata)
• with aerial rootlets with matted pads – form as adventitious roots on stem without
absorption function, such as English ivy (Hedera helix).
Material and energy investments are dedicated to their linear extension growth and
efficient transport system (connecting leaves with roots) over long distances (e.g.
Calamus ratang reach a length of up to 300 m). Stems lack self-supporting
systems and they are dependent on the support plant in reaching the irradiated
area. Sometimes lians are also referred to as skeleton parasites, that is when they
climb as high as the crown of a bearing tree, overshadow it and let the tree gradually
die.

1.1.5 What are the benefits of trees?

A large volume of occupied soil gives them a better chance of getting the necessary
amount of water and mineral nutrients; they can accumulate storage substances
of organic nature, water and nutrients in their bulky bodies so that they could
overcome unfavourable periods for growth. The lignified stems (branches and
trunks) carry reserve buds which can easily replace lost crown parts (damaged by
frost, drought, wind, fire etc.). A vast foliage area is produced very quickly at the
beginning of the growing season, which is an advantage over herbs in the use of
solar radiation for photosynthesis – this also extends the growing season. Reaching
an area of faster airflow and milder temperature fluctuations makes their
reproductive structures better protected; they can also utilize air movement for
propagating pollen and diasporas (fruits, seeds and shoot fragments capable of
rooting, such as crack willow – Salix fragilis). All growth forms of woody plants
support the development of large forests and bushes. Cooperative relations
between individuals (e.g. by root grafts, mutual shading, creating favourable
temperature and humidity conditions, mitigating the damage by wind etc.) together
with competitive relations (competition for water, nutrients, oxygen and symbionts
in the soil, the favourable irradiance above the soil surface) are run in plant societies
(corresponding to certain environmental conditions). In this context trees can reach
ages and sizes close to the species potential optimal. Trees are the most
demanding growth form. In unfavourable conditions many tree species shift to a
shrubby form when they grow old.

1.1.6 Disadvantages of tree growth and possible solutions

The main disadvantages of trees are high demands on supply by photosynthates

that arise in green parts of plants (especially in leaves, but also in petioles and young
stems). These green parts must produce enough energy to cover needs of the
extensive growth of the below- and above-ground systems, the annual creation of
new, functional conductive pathways and mechanical and static stability of the
organism and for securing covering tissues to new and growing tree surfaces.
Photosynthates are also needed for the synthesis of materials, construction of

8
defence structures, forming osmoregulatory substances, symbioses investment
(mycorrhizal fungi) and for making their own storage supplies for survival periods of
dormancy and resuming growth in spring. Adult trees also have increasing demands
for security of generative (sexual) reproduction, i.e. the formation of flowers, fruits
and seeds. In all these conditions foliage of trees works at a much lower level of
hydration than the herbs or low shrubs do. This is because the driving force for
transport from roots to leaves (i.e. water potential difference between these two
points) in trees must be large enough to overcome any resistance that may occur on
the long flow pathway of water from the soil to the atmosphere. The growth and
stability of a tree depends largely on the vulnerability of the "thinnest place" –
stem.
The conductive and mechanical systems of a stem are oversized several fold in
healthy individuals, but material and energy input into the structure of a tree cannot
be reused even in the condition of a critical starvation. Many tree species are able to
reach the "optimal design" in unfavourable conditions, which means to exclude
a stem, recover branches from the stem base, shorten the distance between leaves
and roots, reduce the demands of the water regime, in other words, to shift from the
tree growth to the shrub growth. We can seldom encounter the opposite direction –
rejection of the injured stem, so that the roots grow from the base of the crown (e.g.
under bark or through the hollow in the stem) towards the soil. Thus, the support and
storage functions of the stem are partially replaced and at the same time the
resistance of solutions movement is reduced on the way from soil-plant-atmosphere
system.
Slow wood production together with a reduced speed of the radial growth of
branches (branch overhanging) are other tree strategies leading to efficient energy
balance.

1.1.7 Tree reaction on the unfavourable condition

Tree species as long-living plants do not live only under favourable conditions for
their existence, growth and reproduction. Large energy demands are required by a
tree to overcome unfavourable conditions and stress, which cannot be just escaped
from by trees physically as they are immobile organisms with fixed root systems in a
particular area. The process of disturbance and reaching a new equilibrium
takes place at the level of cell membranes and organelles, and it extends to
individual cells, plant tissues (especially meristematic, photosynthetic and
conductive) and individual organs (e.g., leaves, roots). Finally, it also concerns whole
individuals, i.e. autonomous elements of the ecosystem (whether natural or artificial).
At first, there is a stress stimulus, whereupon a stress response is triggered. By
the term "stressor" we understand the factors of external or internal environment that
cause the organism threat to its functional relations. The stress response is a
change of the functional status (e.g. bent branches for light absorption –
phototropism, diversion of branches and their growth from consistently irritate subject
– tigmotropizmus, remodelling or termination of elongation growth of roots in
compacted soil and restoration of root growth in a more positive direction, etc.).
The outcome of the stress response, which individual tree species are equipped with,
regarding the conditions of the areas of their original extension may include:
a) avoiding the effects of the stressor (e.g. inner leaves, especially auxiliary buds
of woody cushion plants are hidden from temporary drought and heat)
b) adaptation, which means adapting of sensitive structures so that it can withstand

9
the effect of the stressor – (e.g. the sunlight leaves with modified inner structure,
carried on short branches in clusters are more tolerant to repeated insufficient water
supply)
c) resistance to a certain extent of a stressor, i.e. achieving resistance (for example
vegetative buds of conifers and deciduous trees can resist freezing −90 to −196 °C in
the areas with long and tough winters; in the short summer this resistance is
missing).
These examples are known as eustress, which is a stress essential for life, a driving
force of changes leading to the ability to live in a given environment (even
optimal).
The failure of a stress response, i.e. the destabilization or even death of an individual
is caused by one or more conditions, the sum of whose is known as distress. It is the
destructive effect of the impact to a sensitive taxon. Stress response goes through
certain stages. The alarm stage is the pledge of all existing functions, in particular
the synthesis of various agents and growth restriction. Affected tissue respirate
heavily and break the existing structure – catabolism predominates over
anabolism (but spiral shock may spin and cause acute damage and death – such as
peripheral parts of leaves due to salt or bleached districts in immature leaf blades
due to intense solar radiation, etc.)

The restitution stage could follow, resulting in the correction processes of protein
synthesis and protective substances or even replacing the lost parts by a new
growth. Regularly repeated stressors during phylogeny of a species are preceded by
plant precautions, the hardening stage. During this stage the components of cell
membranes are substituted, osmotically active substances are released to free water
to avoid the formation of ice crystals, etc. Trees can achieve resistance by gradual
preparation (see above). Sometimes a chronic injury can be observed in cases
when changes and changing stressors occur irregularly.
A range of stress responses are nonspecific. On the cell level changes in the enzyme
activity (reductases: glutathione, peroxidases), biosynthesis of polyamines and
antioxidants (ascorbic acid and tocopherol), the type and content of the osmoticum
(proline, betaine and polyol), the synthesis of secondary compounds (polyphenols,
anthocyanins, terpenes), synthesis of stress hormones (abscisic acid, jasmonic,
ethylene), changes in the properties of membranes, increased respiration and
decreased photosynthesis. The result of the changes in the relationship between
respiration and photosynthesis (i.e. increasing the proportion of ADP/ATP) is the
depletion of the trees and the lack of promptly available energy. The production of
biomass is reduced (living matter), growth irregularities and reduced fertility occur
(poor internal coordination of growth results in the allometric relation changes
between production, photosynthesizing and consumption parts), and, furthermore, a
tree gets old prematurely (increased necromass – dead matter of crown, stem, roots).
Internal factors that determine the susceptibility or resistance of trees to a given
stressor or complex of stressors, are: a taxon and eco-element (i.e. shady, sunny,
xerophyt, mesophyt etc.), ontogenetic stage, dormant stage or stage of
vegetation activity.

1.1.8 Ontogeny of tree growth

Tree growth, no matter which evolutionary branch it applies to, can be characterised
by the following conditions:

10
a) trees must be mechanically stable (energy to grow and support structures is
provided by leaves),
b) aqueous solutions of the nutrients must be received mainly from the soil and
quickly transported to the leaves,
c) organic materials – products of leaf synthesis – need to be conducted fast
enough and in required quantities to the shoots, flowers, fruits, branches, stems
and roots
d) all parts of tree need to have a sufficient available storage space for water,
nutrients and storage substances.
These characteristics are essentially contradictory as all conditions must be met
and all require energy. Thus, the resulting structure and shape of the plants is
their compromise. This compromise may be reached in the ontogeny, i.e. in the
development of individuals from seedlings to adult trees by these means:
• root system, stem and crown develop simultaneously – such as spruce, beech
– tree appearance is preceded by the initial phase, i.e., roots and leaves develop
while the stem grows in diameter. Only after achieving the required foliage area and
soil occupancy the trunk will elongate (especially ferns, cycads, some angiosperm
species that lack the so-called secondary growth – such as palms),
• a dense root system is formed close to the soil surface, i.e. at the tree base, a
false trunk is formed by intense branching (extinct ferns and present polycormons
palm trees)
• epiphytes living in crowns of host trees, too, may present the initial stage of trees
ontogeny. They let aerial roots down to the soil surface that gradually grow thicker
and stronger, thus forming a stem, which expands in the soil into an independent root
system (some species of the genus Ficus)
Ontogenetic stages can be divided in two periods: the dependence stage – the
period when a young plant uses the reserves stored from the endosperm or
cotyledons (A), and the independence stage – when photosynthesis has become
active (B):
A. heterotrophic period, the individual's life depends on photosynthates supplied by
the parent plant, namely:
1. seed stage and germination
B. autotrophic period, when the new plants can produce and process their own
organic inputs for their growth and development
2. germination (since the full function of the first assimilation leaves)
3. juvenile stage (it is typical for example in thin Norway spruce needles still
undifferentiated into the shady and sunny type)
4. virginal stage (tree habitat is similar to the adult tree, but still does not
flower or give seeds and fruits)
5. stage of younger adulthood (from the first bloom). It should occur after the
achievement of the required range of the stock compartment. In fruit and ornamental
trees it occurs before they reach 10 years; if the vegetative propagation or grafting
was used, it can emerge even much sooner. Domestic forest trees bloom at the age
of several decades, i.e., after an almost final volume of the crown is reached.
A repeated strong stressor, such as undercutting and lateral contraction of roots in
nurseries, could promote early flowering in these trees.
6. stage of middle adulthood (mean maximum crown and root space is
reached and quality seeds are produced)

11
 7. stage of late adulthood (rounding or downsizing top of the crown with
monopodial branched species, reducing the amount of crown, dieback of deep roots
and a gradual transition to flat root system)
8. senescence stage (prevalence of necromass over biomass, reduced
fertility, reduced seed germination)
9. stage of senility (tree survives by a few branches associated with a narrow
strip of cambium with partly living roots)
10. death of the individual
Trees are highly sensitive to stress factors in the younger ontogenetic stages.
However, they adapt most efficiently to their environment just in these stages.
Many seedlings live or at least survive on very unfavourable sites. On the other hand,
the shock from transplanting from a nursery, where seedlings were growing in the
optimal conditions (fertilization, irrigation, soil aeration, etc.) into the environment
dramatically different and unfavourable (compacted soil, changes in temperature,
humidity, light condition) can meet the tree species unprepared. This could lead to
numerous and repeated plantings mortalities. At the end of a tree life (from late
adulthood), its adaptability is reduced and sensitivity to changing and frequent
occurrence of stressors increases.

1.1.9 Dynamic of tree activity

During the long life of trees rhythms and long term cycles of increased and
decreased activity get repeated. The shortest rhythm is daily (circadian) and is
associated with changing light and dark parts of the day. In the light part
photosynthesis should prevail and towards the end of the night, when trees may have
the greatest water content, intense volume growth of cells should occur. Some
sensitive plants demonstrate a monthly, the circa-lunar rhythm, which is related to
the increasing and decreasing moonlight intensity. The activity of most species of our
flora is clearly subject to an annual cycle (cycle circa-annual), i.e. changes due to
the change of seasons – spring, summer, autumn and winter, when the driving factor
is the temperature.
The daily rhythm depends on the proportion of the length of day and night, i.e. on
the photoperiod, whose changes are related to the annual cycle. In our latitudes the
light part of the day extends from the spring equinox to the second half of June with
the longest day and shortest night. Then, the night gradually lengthens and by mid-
July the nights get colder (i.e. change termoperiodicity – temperature difference
between hot day and cold nights). In the second half of September the length of
day and night is equal (autumn equinox) and then the night lengthens until
December. Since the end of December till March the light is on the increase again.
Many of our trees are already at the stage of cold enforced rest (post-dormancy) in
the early spring part of the annual cycle. It depends on the weather conditions
development, especially on the last freezing days in February. If February is warm,
some trees (poplar, birch) would already begin the activity of their roots or flower
buds (hazel). Many trees and shrubs bud and bloom in April, gradually moving to the
stage of full vegetation activity (June, July, exception is for example lilac, which
already declined to an internally controlled dormancy at that time). Shortening the
length of the light part of day, cooling at night and typical weak rainfall result in trees
accumulation of substances that limit cell division and growth, accelerate fruit
ripening and leaf senescence and finally cause falling leaves in the internally
controlled hibernation – deep dormancy. It is not a rule that summer weather is

12
favourable for tree species every year. Plant drying (hydroperiodicity) may occur in
weaker rainy years and at higher temperatures. Trees may respond to drought by
early calming of their growth initiatives, the forced rest, pre-dormancy. In a more
favourable year, it often happens that even species for which such behaviour is
unnatural begins blooming again (Horse Chestnut – Aesculus hippocastanum), for
which a considerable amount of energy is lost. Trees released from deep
dormancy, are driven by internal phytohormonal relations (in particular the level of
abscisic acid – ABA) by the action of frost temperatures and higher insulation (i.e.
condition, which decay ABA). Unlike most deciduous trees, conifers (e.g. Norway
spruce) lack internal control and only have an imposed rest.
Also during the annual cycle periods dangerous to trees may be identified. It is
mostly in spring and autumn, i.e. the transitional period, in which the likelihood of
fluctuating temperatures is high. Late frosts in spring damage young shoots, or
worse, lateral meristems of trunks and branches. In autumn, early frosts may occur
when the trees have not "ripen" yet the late part of the annual rings (completed
lignification of summer wood). The highest sensitivity is in the period of a rapid
growth. Trees are most resistant during their dormancy, a time of minimised
metabolism and energy exchange with the environment. Transpiration flow is
interrupted, water content, movement and metabolism are reduced and growth is
interrupted.

2 Plant Cell

Eukaryotic cell, such as the minimum structural, functional and reproductive

unit, is an open system with the target behaviour of autoregulation and
autoreproduction. This means that it grows, respirates, makes internal movements,
exchanges substance, energy and information with the environment and responds to
them, rebuilds and newly creates its own structure and gives rise to a daughter cell
during the lifetime. This ability depends on:
• its internal memory – inherited and carried out according to the needs;
• the compartmentalization – dividing of its interior by a membrane
• the cytoskeleton, which is involved in the internal movement.
Living cells form a protoplast, consisting of the cytoplasm (cytosol – colloidal
solution of water, protein, fat, saccharides), complex membranes, organelles,
nucleus, vacuoles and ergastic substances produced by cell activities (crystals,
rubber, oil, resin) (Fig. 2). A growing cell builds the cell wall on its surface that may
be part of the inner skeleton of plants (sclerenchyma) or its vascular system
(tracheids, vessels) after the death of protoplast. Not all plant cells build the cell
wall. These include: gametes, sexual cells, i.e. the female (ovule cell) and male
(pollen generative cells) which fertilize together.

13
Fig. 2 Eukaryotic plant cell (adjusted according Procházka et al. 1998)

2.1 Membrane complex, semi-autonomy organelles and cytoskeleton

The membrane complex of the protoplast secures segregation and transport inside
and outside the cell and organelle space, allows for the creation and sustainment
of concentration gradients, electric potential and water potential difference, it
locates synthetic and degradation processes and enlarges reaction areas. The
surface of a life cell body is covered with a simple cytoplasmatic membrane
(plasmalemma), which is semipermeable, semifluid, made of lipid-proteid mosaic. It
is asymmetric. It controls transport of nutrition and metabolites, cell wall structure,
reception of exogenous signals and connection with the neighbouring cells. The
endoplasmatic reticulum is a 3-dimensional matrix of simple membranes that make
flat containers and tubular creations. It divides the cell space, enlarges the inner area
and speeds up the transport of material inside a cell as well as between cells. The
synthesis of protein takes place on the endoplasmatic reticulum surface, while the
synthesis of lipids and saccharides takes place inside. It takes part in the biogenesis
of organelles. Containers of Golgi apparatus are attached to the endoplasmatic
reticulum (ER). The Golgi apparatus work with the output of ER. The processes of
synthesis and transport of cell wall material, secretion activity and creation of
vacuoles are carried out there. Tonoplast is a similar membrane as cytoplasmatic
membrane and it divides cytosols from true water dilutions of vacuoles. Vacuoles are
primary storage spaces of water, saccharides, amino acids, proteins and lipids; they
might be part of lythic (disintegrative) and excretory apparatus and might have
protective (bind dangerous ions), defence and alarming functions (toxins, pigments).
Ergastic substances include starch, lipids, proteins as storage substances,
cellulose, lignin as structural substances and substances with physiological and
ecological functions (crystalline inclusions, aromatic substances etc.).
The plant cells contain two types of semi-autonomy organelles (plastids and
mitochondria); their structure are similar to simple prokaryotic cells. In the
prehistory they used to be enclosed in eukaryotic plant cells as endosymbionts.
Chloroplasts (centres of photosynthesis) surrounded by a double membrane are a

14
good example of these. Their interior space is divided by thylakoids into the grana,
as well as thylakoids that connect grana (Fig. 3). In the chloroplast membranes,
there are pigment-protein complexes needed for the reception of radiant energy and
its transformation into the energy of chemical bonds of saccharides. Mitochondria
(centres of respiration) are also surrounded by a double membrane. Their interior
surface carries elements of respiration chains and enzymes needed for the formation
of ATP (adenosine triphosphate), there is a cycle of acid citric, NAD (nikotinamid
adenin dinucleotide) reduction processes and the degradation of fatty acids. What
decides on the semi-autonomy of these organelles is their own spiral DNA
molecule (deoxyribonucleic acid) and the related proteosynthesis.
The cell inner structure, organelle position, exterior shape of the cell and its directed
growth are guided by the cytoskeleton whose specific proteins are organised into
microtubules, microfilaments and short microtrabecular fibres. It carries out
transport of cytoplasm using energy (from ATP), controls the position of chloroplasts
towards the radiant flow, chromosomes during cell division and it also decides on the
place and direction of cellular microfibrils in cell walls.

Fig. 3 Chloroplast. Left – oval shape of plastids in the mesophyll cells of spruce
needle; middle – picture of inner plastid structure from electron microscope; right –
schematic picture of inner plastid structure (adjusted according Solomon 1996)

2.2 Nucleus

Nucleus is the centre of the interior memory which is sustained, multiplied and
transported into the actual cell and even from generation to generation (of cells as
well as organisms). It is separated from the cytoplasm by a double membrane with
pores interconnected with the endoplasmatic reticulum. It contains a nucleolus,
nucleous matrix with ribosomes, nucleoproteins and chromatin, i.e. nucleic
acids that function as a genetic apparatus.
These are heterobiopolymers based on the specific sequence of nucleotides
with purine base (adenine, guanine) or pyrimidine (cytosine, thymine in DNA and
cytosine, uracil in RNA). These molecules also contain saccharide pentose
(deoxyribose in DNA, ribose in RNA) and a phosphate group. DNA has a shape of a
double helix, opposite bases of both the fibres (connected only by a hydrogen bond)
are equal (adenine opposite thymine, guanine opposite cytosine). The specification
of these acids is given by the sequence of nucleotides, whose triplets encode a
specific amino acid, and the sequence of triplets is given by the sequence of
amino acids in proteins. New DNA molecules are always copies of already existing

15
ones. After the double-helix molecule is unwound and its two fibres are separated,
new copies of these fibres are synthesised, so that two new molecules were made
from the original one (replication, reduplication).
Proteins are heteropolymers as they are made of the selection of more than 20
amino acids in various sequence, various number of repetition of individual
passages, various lengths of the chain and various possibilities of spacial
arrangement. Taking into account all these variations, they have different
characteristics (as building blocks and parts of enzymes) and this makes them the
primary source of the variability of life forms and originality of each individual. The
synthesis of proteins proceeds in several stages. In the nucleolus, according to the
model – unwound DNA passage, the sequence code of bases is transcript into three
types of RNA (mediator, ribosomal and transfer) – i.e. transcription. Then these
molecules undergo transformations (mutation – cutting out extra passages and
making new connections) and then they move from the nucleolus to the cytoplasm.
Double-body ribosomes with rRNA (ribosomes aggregation makes polyribosome) join
the mRNA fibre, and when moving along the fibre, they match with individual
activated amino acids, transferred from the tRNA cytoplasm, according to the triplet
sequence – several equal molecules of individual protein are generally synthesised in
this way. They originate by the translation of information from DNA. These proteins
are placed into ER containers, where they are checked and transported into the
places of need, or placed into the Golgi apparatus for further syntheses.

2.3 Cell ontogenesis

The individual cell genesis (ontogenesis) is divided into the division meristematic
stage (embryonic), the elongation stage and the differentiation stage (towards a
specific function). Cell division (i.e. cytokinesis) is followed by the division of its
nucleus – karyokinesis. Nucleous DNA condenses, spiralizes and connects with
single protein histones to form chromosomes. Nucleus membrane gets dissolved,
chromosomes undergo longitudinal division and half of each moves to the daughter
nuclei (due to cytoskeleton), i.e. nucleus equation division – mitosis. In the newly
formed nuclei, transported DNA molecules (chromosomes) are resolved into the
functional simple fibre structure and the nucleous structure is renewed. The amount
of DNA in daughter cells rises to the original value by means of replication.
Cytoplasm as well as all other cell structures, including cell membrane, are multiplied,
and the cells prepare for further replication (meristem cells) or shift towards the
next ontogenetic stage.
Healthy cells (formed by the mitotic cell division) have an even number (diploid) of
chromosomes (i.e. each chromosome type is represented twice there, i.e. in case of
yew tree, the cell nuclei comprise of 4 chromosomes, 12 in spruce, larch and pine,
and 26 in maple).
The gametes genesis – male and female sexual cells – is preceded by another
karyokinesis type – meiosis. It leads to the creation of cell nuclei with a basic set of
chromosomes (haploid) which are greatly transformed by the mutual replacement of
passages between pairs of mutually corresponding homologue chromosomes. A new
individual initiates from the fusion of a male gamete nucleus (generative nucleus
of pollen with a base chromosome set) with a female gamete nucleus (i.e. ovum
cell nucleus which also comprises of a base number of chromosomes). This creates
zygote – the first cell of a new individual – having a double set of chromosomes, one
of whose comes from the mother and the other from the father organism . In

16
angiospermous plants, there is an extra fusion of the second nucleus of pollen
(having a basic number of chromosomes) with the central nucleus of the embryo sac
which is diploid. This is how a triploid endosperm is made. It feeds the plant embryo
until it matures.
The nucleus division, or fusion of nuclei, is followed by the cell division
(cytokinesis). Regarding cytoplasm and its structures, the division is not even.
Some elements reach the daughter cell transformed (e.g. plastids, virus element
etc.), so daughter and mother cells are not equal (this causes e.g. unevenly
coloured stripes or segments of leaves). For instance, it is the prime division of
zygote that really decides which of the daughter cells shall become the basis for the
above-ground and root structures. Onwards, root cells censure a great deal of
information, e.g. that could be used in photosynthesis or blooming. Similarly, mother
cells of xylem and phloem are distinct, though they were initiated in the same cambial
initial. In theory, all plant cells include all information needed to create a new
plant of a given species – this allows for the vegetative reproduction: a sprout
with encoded information on how to build and form shoots develops on a root, or
roots may as well be formed on shoot. In general, in the embryonic growth stage,
there are intense syntheses of nucleous acids, lipids, proteins. The plant needs a lot
of nitrogen and phosphor. The growth regulators of the greatest influence involve
cytokines and gibberellins.
The above described stage of the cell division growth is closed by the construction
of a new partition – cell wall. Its first layer consists of proteins, mainly pectates
(salts of pectate acids). It is called middle lamella or intercellular matter. After that
the new cells concentrate vacuoles, they get filled with water – their pressure from
the inside of the cell wall rises and they grow in size (length of cell may increase of
up to 20 to 50 times). The construction of an extending wall is finalized by the
primary lamella. Cellulose macromolecules (homobiopolymers of glucose residues)
are inserted into the lamella and get arranged in crystalline microfibrils. Microfibrils
may be more or less well structured, interconnected by pectates and hemicellulose
(besides glucose they contain e.g. saccharide xylose) with lower proportion of
protein. The new matter is inserted by means of intussusceptions – the wall grows in
size and does not prevent the volume growth of protoplast. However, bonds between
wall parts are weak, cell wall is not solid and behaves as fluid crystal, and this makes
the cell susceptible to further changes in shape. After the shape has been stabilized,
the ontogenetic stage of the volume growth is closed. What ensues from the
description of this stage is that a plant mainly needs water as it is the force that
extends growing cells and develops a hydraulic pressure on the neighbourhood.
Another needed matter is organically bound carbon (saccharides) widely used for
the formation of cell walls. Regarding growth substances, gibberellins are the ones
that are most important at this stage.
As cells enter the differentiation growth stage, depending on their future specific
function, they build secondary layered lamellae to strengthen the cell wall. The
lamella is formed by the apposition of further layers from the inside of a cell; i.e. the
middle layer is formed first, the primary one follows and the secondary is formed last.
The building substances include: cellulose, hemicellulose, lignin, suberin, cutin,
waxes, silicon dioxide, carbonates, sporopolenines etc. Since this is a centripetal
growth, i.e. from the periphery towards the centre (the reverse process, centrifugal
growth occurs in cell wall formation of pollen), the protoplast area might be
diminished to such an extent that a cell could not survive. In most cases, particularly
in skeleton cells or xylem conductive cells (with variable pressure conditions from

17
overpressure to underpressure as low as 3-4 MPa) the secondary lamella is
formed unevenly. Precise direction and laying construction is driven by cytoskeleton
and complex structures is form. Pits, e.g. simple or branched ones, mediate
connection and communication through plasmodesmata (cell wall channels lined with
cytoplasmatic membrane that connect the protoplasts of adjacent cells) among cells
as well as organs (by means of long and medium distance diffusion), which helps
sustain balance in an individual. Bordered pits (in gymnospermous tracheid walls)
and even perforations (e.g. in vessels, laticifer ducts) enable material flow on long
distances. Various types of thickening (round, spiral, step, matrix) prevent from cell
deformations (e.g. phloem and xylem fibres, sklereid etc.). Construction and
material structure of cell walls reflect their basic characteristics; i.e. insolubility in
water, anisotropy (e.g. birefringence in polarized light and spacial distinct changes in
volume during growth) but also the ability to communicate with the neighbouring
cells or water and gas permeability. There are numerous instances when the cell
wall features are not compatible with protoplast existence, so after the wall is
formed, protoplast dies out and its functions (mainly mechanic, conductive) are
taken over by the cell wall.
There are two types of factors on the differentiation stage of cell life: endogenous –
i.e. genetic, enzymatic, phytohormonal activities, and exogenous – e.g. climatic, soil,
topographic (relief and morphology) and biotic (e.g. galls on leaves brought about by
fault cell differentiation as an impact of insect activity). There is an increase in the
consumption of mineral sources and the demand on the optimal photosynthesis and
respiration. Some kinds of already specialized cells are capable of dedifferentiation,
i.e. return to the embryonic stage and could restart division. This is why surface
injuries may regenerate and heal.

3 Tissues

Woody species have true tissues (originated from a single cell) and are defined as
groups of cells of the same shape and functions. They share some common
features. For instance, the cell walls are interconnected by intracellular material
(middle lamellae comprised of calcium magnesium pectates). Life cell (protoplasts)
are interconnected by plasmodesmata with the tubular endoplasmatic reticulum.
By means of these, all life cells of the whole system form one unit – a symplast. It
can actively transport (by speeded up diffusion) substances that have been formed or
transported through cytoplasmatic membranes in another part of a plant.
Another feature of tissues is the presence of intracellular spaces. They are formed
in three different ways:
- schizogenous – by splitting of middle lamellae (e.g. in corners of cells or
between guard cells of epidermis) or by tissue injury when intracellular material and
walls may be dissolved (Fig. 4)
- rhexigenous – by tearing off tissues due to fast growth of neighbouring
tissues (e.g. in pith, especially in monocotyledonous liane stems)
- lyzigenous – by dissolving of cells (essence reservoirs isolated by lignified or
corked walls of neighbouring cells)

18
Fig. 4 Schizogenous intercellular space – Resin duct in the pine needle developed by
splitting of middle lamellae of secretory cells.

Intercellulars hold a substantial role in transport of gases and water (e.g. in leaf
spongy parenchyma, leaf hydathodes and lenticel filling tissue) and in concentration
and isolation of protective substances (e.g. resin canals and silicae containers).
Schizogenous intercellulars with aeration functions grow at decreased aeration in
inner tissues (in roots during soil flooding, in stems and branches lay on the ground
etc.) They give such a tissue its name – aerenchyma.
Intercellular areas (except for excretion glandule, resin and laticifer ducts) as well as
interior areas of dead xylem cells (e.g. vascular bundles) create an apoplastic
matrix – a communication system which transfers matters in gaseous, liquid and
solid phases without crossing barriers of cell membranes. Even saproparasite
organisms could spread through the apoplast.
Idioblasts are commonly present in tissues – these are cells that considerably
differ from the majority of cells by the shape, type of the cell wall (e.g. sclerid) or
by the contents (crystals, tannin in vacuole, e.g. tannin cells near phloem in spruce
etc.)
Tissues are categorised by various criteria. From the perspective of a cell wall
type, cell lifespan and function these tissues are distinguished (Fig. 5):
- parenchyma – thin-walled cells, mainly living, capable of division, metabolic,
storage etc. functions
- collenchyma – living cells with unevenly thickened cell walls. If they thicken at
celll corners this is called angular collenchyma, if thickened walls are parallel to the
organ surface, this is called tangential collenchyma, if it contains intercellulars, it is
known as lacunar collenchyma); it acts as a mechanical support, metabolic support, it
can renew division and give rise to healing tissue
- sclerenchyma – tissues that contain mainly dead cells whose function is
taken over by strong lignified walls. The walls are filled with various types of
narrowings as well as thickenings – round or spiral ones that do not slow down the
elongation growth of neighbouring cells or matrix and staircase thickening that occur
in the parts of a plant where the elongation growth is no more as active as it used to

19
be. Sclerenchyma cells have mechanical (supporting) and transportation functions.
They transport solution from roots to the above ground parts.

Fig. 5 Tissues division from the perspective of a cell wall type and cell lifespan:
parenchyma, collenchyma and sclerenchyma

There are two types of cell from the perspective of cell shape – isodiametric (i.e.
star-shaped, spongy, pallisade etc.) and prosenchymatic (highly elongated cells).
From the perspective of the rate of differentiation, the following types are defined:
meristem – dividing, embryonic, which never reaches growth and differentiation
stages, localized in growth zones as apical primary meristems (on stem, bud and
root tips, or on a growing section of a leaf) and lateral secondary meristems. By
cell differentiating, meristem gives rise to permanent tissues, also divided into
primary and secondary ones. They are functionally arranged into the systems of
covering, vascular and basic tissues.

3.1 The origin of primary meristem

The whole tree organism originates in a single cell called zygote. It is the initial cell
of an offspring organism which was formed by the fusion of a sperm cell (male) and
an ovum (female) cell, called gamete.
Zygote is bipolar, i.e. there already exists a physiological difference which later,
when organs and whole organisms are formed, directs the flow of material,
photosynthates, hormones and other controlling substances. This polarity also
directs the formation of adventitious roots and buds from stem and root. In case of
stem, new roots are formed at basal (proximal) cutting area and new buds at apical
(distal) poles; it is opposite in root sections.
The zygote polarity defines the basic features of root and above ground systems
as early as the first cell division. The onward cell dividing with genetic regulation
leads to the creation of an embryo of a plant. Embryo contains all merismatic

20
cells, i.e. cells capable of intense dividing. A mature embryo contains: a radicula –
later develops into the primary root; a hypocotyl – primary part below cotyledons,
stem part bearing primary leaves – cotyledons (monocotyledonous plants have one,
dicotyledonous have two, gymnospermous have multiple cotyledons). In the tray of
cotyledons, there are primordia of the first lateral buds and epicotyls between
cotyledons – a primary above cotyledon part of stem enclosed by a plumula, i.e.
apical shoot of the above ground system (Fig. 6).

Fig. 6 Schematic picture of germinate plant. Left – gymnosperm; right – angiosperm

3.2 Primary meristems

All cells of a plant divide during the germination and growth, but they gradually
lose dividing capability. They start growing towards the base (in proximal direction)
and focus on special functions in dependence on their positions. This means that
meristems move in distal directions, i.e. above-ground grows towards the sunlight
and to the root apex (grow in direction of gravity).
Dividing character is retained only in the apical cell groups – initials, that occur
at the main root apex, later in lateral roots and on the stem apex and lateral shoots
(Fig. 7 and 8). The initials are directly connected to the embryo cells
(archimeristems); and together with the newly divided or just being divided derivates
they form a protomeristem. Its older derivates still capable of dividing form a
protoderm (giving rise to the primary epidermis tissues), procambium (whose
dividing cells arranged in stripes differentiate into primary vascular tissues – xylem,
and phloem) and the basic meristem (predecessor of epidermis, pith, pith rays in
stems and roots, mesophyll in leaves).
Protomeristem, protoderm, procambium and basic meristem are primary apical
meristems responsible for the elongation growth and for the formation of a set of
primary permanent tissues; their spacial arrangement decides on the primary
structure of organs. The meristems may retain in the latent state (suppressed
buds) for a long time. They generate leaves and buds in their trays as basics of
lateral branches. Suppressed buds are a storage which can easily regenerate a
plant. On the other hand, they can lead to the formation of various tumours by

21
pathogens. Between stem nods with leaves from the primary meristems, there is an
intercalary meristem which allows for later elongation of internodes. The shape of
leaves ensues from different activity of the residue meristems on the basis, tip and
edge of their blades.

Fig. 7 Primary meristems of shoot apex. Leaf primordia protect apical meristem.
Axillary buds are form in the leaf primordia base.

Fig. 8 Primary meristem of root apex. Root cap protect root apical meristem and
enable root growth in the soil. Lateral root are form later.

The primary structure of stems and roots is typical to some herbs (e.g. in
Ranunculaceae family) and monocotyledonous woods that do not secondarily
grow in diameter. The structure of the other woods that form secondary
meristems is more complicated. Its appearance can be well described by means of
three mutually perpendicular sections. The cross-section is perpendicular to the
longitudinal axis of an organ. The radial section is longitudinal and crosses the

22
centre of an organ. The tangential section is longitudinal too, but it does not reach
the centre, it remains at the periphery.

3.3. Permanent primary tissues

3.3.1 A system of primary surface tissues

The differentiation of protoderm derivates in above ground organs leads to the

formation of compact, unwettable epidermis covered with cuticle and epicuticle
wax from the outside. If epidermis is multilevel, then the inner layer is call
hypodermis. It is gas proof except for stomata, it protects the inner tissues from UV
radiation which effectively absorb, and it serves as a water reservoir. By means of
opening and closing stomata, it protects a plant from withering, overheating by
transpiration (distribution of water gas to air) and from biotic factors. Trichomes
(outgrowths of epidermis cells) are helping to fulfil epidermis functions. These
trichomes might be single-cellular or multicellular (Fig. 9), branched or unbranched,
glandular. More massive formations are called emergencies which consist not only
of epidermis but also inner tissues (Fig. 10). These involve: prickles (rose,
raspberry), glandules (in ash tree or chestnut shoots), or pollen section in flowers.

Fig. 9 Multicellular trichomes of Viburnum rhytidophyllum

23
Fig. 10 Prickles (emergencies) of rose. They consist not only of epidermis but also
inner tissues. Unlike spines they are form irregularly.

In case of roots, rhisodermis is formed. It has weak cell walls, it is less water and
gas resistant, it swell when wet, it has no cuticles or stomata. The rhizodermis
produces acids and exoenzymes that disturb the substrate and nutrients. Vacuoles
contain osmosis active solutions, cells lack chloroplasts, stomata, cutin and wax,
but they have leucoplasts. There are rhizomes, parallel to trichomes, but unlike
those, they have the absorption function. They are shorter and less condensed in
trees than in herbs, they are lacking in case of root colonisation by mycorrhytic fungi.
Taking into view the characteristics of rhisodermis, roots are very sensitive to drying
out. Fine roots of seed plants removed from the soil, left in the sunshine and on dry
air may die in a couple of minutes.

3.3.2 A system of primary basic tissues

Primary cork called cortex is a basic multicellular tissue between the epidermis
and vascular tissues. In roots, outer layer (exodermis) cells have protective
functions (corked later replaced the rhizodermis), medium layer (mesoderm) have
transportation and storage functions. Endodermis, inner layer is made by cells with
widened, corked stripe of cell wall (i.e. Casparian strip) and cells above vascular
tissue that are permeable – this is a means of endodermis control intake of
substances into vascular tissues in the central cylinder. Primary cork of shoots has
a photosynthesis function, storage function (life parenchyma cells) and a
mechanical function (collenchyma and sclerenchyma). Life primary cork cells are
capable of remeristemation after an injury.
The primary cork is linked to the central stem and roots parts by the pith rays. They
secure horizontal transport of substances, e.g. oxygen from the atmosphere and
saccharides from xylem to dividing and growing cells in the cambial zone. Pith ray
cells are a reservoir of crystals, phenolic and toxic substances; there are horizontal
resin canals in many gymnospermous plants. Pith occurs inside stems, hardly ever
inside roots. Its thin walled cells in young stems may contain chloroplasts or starch,
later on secretions and lignin in cell walls.

24
Basic tissues involve mesophyll in leaves – a tissue with photosynthetic functions. It
is the most significant producer – source of saccharides, molecules rich in energy
adenosintriphosphate (ATP). These are formed using solar energy for the chemical
transformation of atmospheric carbon dioxide and water.
Basic tissues have a lot of functions: assimilation, reservoir, mechanical
(collenchyma, sclerenchyma) and excretive (hydatods, nectarines, resin ducts etc.).

3.3.3 A system of vascular tissues

It is a vascular bundle that interconnect distant parts of a plant by means of

distributing mineral substances, saccharides, regulation molecules. Vascular
bundles may be complete (containing both xylem and phloem parts), or one of
these parts may be absent (incomplete vascular bundles, e.g. apical parts of tree
tops or foliage mesophyll). Their primary structure is retained in
monocotyledonous woods and woody ferns and their arrangement is scattered
and vascular bundles are either concentric (phloem-centric, phloem is surrounded
by xylem, e.g. Pteridophytes) or enclosed collateral (centripetal xylem and
centrifugal phloem – palm trees). Such vascular bundles could be divided in two
subgroups at the final stage of the development – older protoxylem (thinner
vessels, cell walls are reinforced by circle or spiral thickening) and protophloem with
obliterate or dilatate (i.e. non-functional) cells. The latter is younger – with fully
developed and more efficient younger metaxylem with broad tracheids or vessels
and metaphloem with functional sieve tube and sieve cells. The mentioned vascular
bundles are created by general and complete differentiation of procambium strips.
Vascular bundles in stems of gymnospermous and dicotyledonous woody
plants are collateral, open (i.e. procambium stripes are only partially differentiated,
so cambium between the phloem and xylem remains active), arranged in circle,
separated by rays. Protoxylem lies on the periphery of pith and metaxylem is located
centrifugally . As early as the first year of stem development, the metaxylem
neighbours with the secondary xylem (deuteroxylem) in stems and roots. The
passage is gradual, it may even be less perceptible. On the edge of the central
cylinder, there is protophloem which is no more functional, centripetal metaphloem
and towards cambium there is completed secondary phloem (deuterophloem).
In early stages of development, roots have radial vessel system, whose individual
incomplete vascular bundles rays where xylem and phloem groups are
alternating. Vessels in monocotyledonous roots are organized in multiple groups.
Gymnospermous and dicotyledonous plants have usually monarch (one group of
xylem and phloem), diarch (two groups of xylem and floem), triarch (three groups of
xylem and phloem) fine roots. Cambium in gymnospermous and dicotyledonous roots
start to form below the phloem groups and above xylem groups later on. The radial
vessel bundle passes into multiple open collateral vessel bundles arranged in a circle
and separated by phloem-xylem rays.

3.4 Secondary meristems

3.4.1 Cambium

Primary structure in woody plants is retained only for a short time. Procambium is
active and changes into the secondary lateral meristem – cambium. Below the
outer layers of tissues (epidermis, primary cork, primary phloem), cambial initials

25
form a coherent cover of a cylinder that appears as a cambial circle on the cross
section. It contains sections of fascicular cambium responsible for extending of
vessel bundles, and sections of interfascicular cambium giving rise to phloem-
xylem rays. Bundle cambial initials are longitudinal in direction of a long axis of an
organ and separate mother cells of the secondary xylem centripetally, and mother
cells of the secondary phloem in the opposite direction. Tranverse partitions
between daughter cells are parallel with the surface of organs (periclines). When
daughter cells arranged in radial lines grow, the whole organs grow with them. As
early as the first year of activity, some initials of cambium go under such a division
that cell walls separating daughter cells are perpendicular to the surface of the organ
(anticlines). It results in the creation of new initials of interbundle cambium, i.e. in
the rise in diameter of the cambial circle and rise in number of vessel bundles and
rays. In that case, rays do not reach as far as the pith or primary cork. Such rays are
called secondary phloem-xylem rays.
Intrabundle cambial initials are nearly isodiametric (same size in all directions). They
divide ray cells of the same type in both directions – centripetally as well as
centrifugally. Cambium is responsible for diameter growth of the stem and root
systems. However, while it creates one annual ring (early and late wood) in the stem
system during a growing season, and the number of annual rings more or less
corresponds to the age of a tree part, in case of roots, it is different. There are more
growth waves in a year. These are irregular and their boundaries are less obvious, so
it is not that easy to determine their age. In evergreens, cambium appears in leaves,
too (in veins of the first or second order) and there are more phloem than xylem
layers in needles.
Phloem is conductive tissues that transport carbohydrates from the locations of
syntheses to the consumption locations: meristems, root systems, flowers, fruit or
buds). In gymnosperms phloem tissue is made of short-life sieve cells with dense
sieve areas located mainly in cell walls with connection to albuminous cells, which
neighbour with pith rays. In angiospermous species, this tissue is formed by
elements of sieve tubes (Fig. 11). The tubes are made of cells arranged in vertical
lines. These are longer conductive passages. Sieve tube elements are elongated in
direction of a long axis of an organ and they dissolve cell nuclei during the process of
differentiation. This means that they lose the information bank, cannot react to
changes that may occur and their lifetime is extremely shortened. Longitudinal cell
walls are perforated into the sieve area similar to sieve cells, but there are large
sieve pores on the connection of two sieve tubes. Cytoplasmatic membrane passes
from one element to another and distributes organic material (esp. carbohydrates)
into all places of consumption in a plant body.
The content of sieve tubes demonstrates positive pressure (approx. 3 MPa) on the
inner walls, carbohydrate solutions may be concentrated even more than 20 % and
the speed of transport is approx. 1 m . h-1. The sieve pores are plugged by
polysaccharides callosa originated from glucose remains, which is produced and
stored at the end of the growing season. This process close conductive pathways
in sieves sooner than usual. Sieve tubes may be accompanied by phloem
parenchyma (soft phloem) and sclerenchyma fibres (hard phloem). Non-
strengthened walls of older sieve tubes allow for the obliteration by the pressure of
the surrounding tissues. The pathways are hardly ever active for more than one
growing season (e.g. in lime or Tsuga canadensis – hemlock). The secondary
phloem is formed by cells divided by cambial initials within open vessel bundles of
gymnospermous and dicotyledonous woods. It is renewed each year. In

26
monocotyledonous woods, e.g. phloem-centred draceana, it can also stay active
without a renewal.
In autumn after the finished elongation and diameter growth of stem in wood parts of
vessel bundles, we could observe protoxylem cells located between typical wood
and pith – they are tiny, thickwalled. Out of the protoxylem, xylem cells are larger in
diameter (metaxylem) and finally, secondary xylem cells are observed (firstly early
wood and then late wood). Small cells from the previous growing season are covered
with a new layer of early wood of large diameter cells and thinner cell walls. This
makes the annual ring border well observable.
If the growth is not interrupted (esp. in young trees), the number of annual rings on
a stem base tell us the tree age. Based on the appearance and arrangement of early
and late wood cells, deciduous trees are categorized into ring porose, semi-ring
porose and diffuse porose ones. A ring of extremely thick vessels which quickly
pass into a number of radial lines of thinner vessels and other wood elements with
smaller diameters (e.g. ash tree, acacia) is formed in spring in trees with ring pores
arrangement. Regarding semi-ring and diffuse porose ones, the inner diameter of
vessels is diminished gradually and the boundaries of annual rings may not be well
observable. The cambial ring grows in diameter as the number of initials multiplies.
The number of vessel bundles and rays slowly grows. The rays that were formed at
the plant origin, stretching from the pith to the primary cork, are called primary, and
the ones that were formed later, are known as secondary or phloem-xylem rays.
These are shorter in the radial direction.
According to xylem type we distinguish between homoxylic (gymnospermous) and
heteroxylic woods (angiospermous). Wood structure in homoxylic is simpler as
there is no division between conductive and specialized cells with mechanical
functions. Both conductive and mechanical functions are controlled by
tracheids; elongated, prosenchyma cells without cell wall perforations. Water
dilutions pass from a lower tracheid to an above one through border pits on the radial
walls. Parenchyma is mainly found in rays of various width and height. Annual ring
boundaries, specifically latewood with thick lignified tracheids of little diameter, build
a barrier from freeze, fungi or insects damage. The essential protective system is
a three dimensional net of resin canals, located in wood vertically, and in pith rays
horizontally. The heteroxylic species (angiospermous, e.g. deciduous trees) contain
special conductive cells – vessels, but tracheids are also present.
Vessels are columns of dead tubular cells with perforated transverse cell walls and
longitudinal walls with pits and thickenings (Fig. 11). In case of ring porose species,
early vessels have large diameter (up to 0.5 mm) and high sap flow (over 60 m . h-1
in lianas). They are often found under a great negative pressure, which occurs when
the evaporation from leaves is faster than the root uptake of water from the soil. In
this case, they are sensitive to functional disorders due to the penetration of air (air
seeding) into the vessel (embolism) or penetration of xylem parenchyma from the
neighbourhood. Parenchyma spreads in the vessels interior, fills the vessel in and
gives rise to thyloses. Xylem fibres (libriform fibres) are responsible for the
mechanical resistance of wood. They are thin, long, usually dead cells with thick
lignified walls. It is mainly parenchyma that is responsible for the storage function.
There are two type of parenchyma - axial, parallel to the longitudinal axis of an organ
and usually accompanying vessels and horizontal, i.e. parenchyma of pith rays.

27
Fig. 11 Vascular bundle is composed of xylem (left) and phloem (right). Both parts
are shown on the cross and radial section. Sieve tube development is shown.

3.4.2 Phellogen

Together with the cambium activity and every new growth of xylem, phloem and
parenchyma of rays, the plant diameter extend and develop a pressure on the
outer layers of cells in the radial direction. This results in the formation of another
lateral secondary meristem – phellogen, which might be located bellow epidermis, in
primary cork or in pericycle in gymnospermous species.
Double-sided dividing of daughter cells is typical of the phellogen. The inner
daughter cells live longer and form so called phelloderm. The outer daughter cells
store suberin in the cell walls; and as they become cork, water and gas resistant,
they gradually die. They function as a dead tissue – cork (phellem) that protects the
inner parts of a tree from the exterior danger. The layer of phellem, phellogen and
phelloderm is considered a secondary cork.
Though, such a system of surface tissues in not permeable. Other elements must be
formed to let oxygen pass to life meristem cells (cambium, phellogen) and life
phloem, xylem and ray parenchyma cells in the secondary cork (or even closer).
These elements are called lenticels (Fig. 12). They are tiny, they have a rounded or
line shaped cracks in the bark of trunks or branches, or in the epidermis of green
stems (e.g. spindle tree) or epidermis of fruits. They originate by quick dividing of
parenchyma cells by the phellogen cells towards the outer parts of stems. The
tissues with rich intracellular spaces aerate neighbouring deeper tissues. The
lenticels are frequently formed above xylem rays. In pre winter period, the lenticels
get covered with water and gas resistant cork cells, thus a protective closing layer is
made. This layer only lasts till spring when it gets torn by the pressure of the newly
produced cells.

28
Fig. 12 Lenticel – cross-section of Sambucus nigra shoot. Faster production of
phellem resulted in epidermis rupture and lenticel formation.

The activity and lifetime of phellogen greatly varies from species to species. In
some species (beech, hornbeam), phellogen cells are able to grow in number by
means of anticline division. The phellogen ring growth is influenced by trunk
thickening, which is carried out by the activity of cambium. Bark of such trees is
smooth. In other instances, phellogen cells are not able to form anticlines and divide
only by means of periclines. The phellogen gets crack and gradually dies out as a
result of an increasing radial pressure by newly developed annual rings. In deeper
primary cork tissues, later on even in life phloem, new phellogen is formed. They
give rise to new layers of periderm under dead outer layers. The repetition of dying
and the formation of new phellogen is a unique complex where dead layers of
primary cork meet layers of periderm, and, later on, dead layers of phloem meet
deeper and deeper formed periderm. This tertial complex of cover tissues is called
rhytidome or bark. When it is peeled off, the older non-functional layers of phloem
are peeled, too. This could be macroscopically observed as a coarse, deeply cracked
bark, which varies according to species (trunk base of old pines, birches etc.). The
following phellogens could occur in concentrated rings – in such a case the bark is
known as ring bark and peels in longitudinal or cross stripes (juniper, cherry), or the
new phellogen is formed deeper and deeper as smaller or bigger lamellas, and
bark peels off as large scales (plane trees) or small scales (spruce) (Fig. 13).

29
Fig. 13 According the activity and lifetime of phellogen great variability of bark are
formed. A – ring bark (Juniperus sp.); B – scale bark (Picea abies)

Another latent meristem is called pericycle. It occurs at the edge of the central
cylinder and together with cambium it may participate in the formation and growing of
lateral roots or formation of adventitious buds or roots, but it can also sclarify and
make a heavy mechanical support of phloem (Aristolochia durior).

4 Roots

4.1 The origin of root

The oldest vascular plants used to be rootless. Even these days, all vascular spore-
bearing plants only have lateral roots of the adventitious origin. Typical roots can be
only found in seed plants. They are developed already in the embryo stage
(radicula).
In gymnospermous and dicotyledonous plants root growth (during germination) is
direct continuation of radicula. Such a root is called primary. Majority of lateral
roots on this primary root are formed as late as the post-embryonic stage. The
outcome is the formation of the allorhitic root system, which is divided in the
primary root and lateral roots. Later on, new roots may grow on old roots, trunks or
branches (leaves). They are called adventitious roots. In monocotyledonous
plants, there is a homorhyzic root system. A large number of lateral roots are
found even in the embryonic stage and radicula dies out in early stages of a plant
growth. A couple of solid roots are based on hypocotyl. Moreover, the root system is
further extended by branching as well as establishing further roots on hypocotyl and
higher positioned stem nodes.

30
4.1.1 General characteristics of root systems in trees

Root is prevailingly an belowground organ, which is heterotrophic and non-

segmental. Along with the development of the above ground system, the root system
grows by means of branching of the primary root. Roots have a radial symmetry and
are bipolar:
- the first pole – root apex, is distal, i.e. distant from the mother root or stem
- the second pole – root base, proximal, i.e. is connected with the mother root
and with the stem base).

This polarity is retained even at injured roots and regenerating root sections: roots
(more numerous and finer) are formed at the distal area and formation of adventitious
shoot buds at proximal area. Root functions depend on photosynthates (esp. on
carbohydrates) derive from the above ground photosynthesising organs (leaves, i.e.
from primary sources), or from cotyledons or endosperm (i.e. from secondary
sources).
Endogenous and irregular branching gives origin to the multilevel root system.
Root branches are called lateral roots of the primary, secondary etc. orders. They are
formed in an acropetal order, in other words, the youngest branches are always
nearest to the apex tip. This type of branching is called monopodial (Fig. 14).
Sympodial branching (lateral branches are more dominant than the primary root) is
less common. This branching pattern can be found in some trees with mykorrhizza
(Pinus).

Fig 14 Fine roots on Norway spruce growing in the aerated and nutrient rich soil
layer. Root branching is monopodial, regular and rich with many active root tips.

31
Fig. 15 Fine root of Norway spruce growing in the heavy and wet soil in the 1 m
depth. Root branching is irregular, mostly root are dying, lateral root are short with
spare root tips.

Lateral branching of roots usually occurs in an endogenous way. The lateral roots are
formed inside the branching root from the latent meristem, pericycle. Cytoplasm is
accumulated at some places in pericycle cells and the cells begin to divide both in the
radial and tangential directions. The base of a lateral root is established. Further on,
the root penetrates the primary cork and rhizodermis to get to the surface. The origin
of lateral roots is not accidental. If the branching pattern is observed in detail, it is
clear that they are arranged in longitudinal lines, i.e. orthostichies. Cross sections
demonstrate that the position of lateral roots is related to the position of xylem and
phloem section of the vascular bundle. In some species, lateral roots originate in
longitudinal lines opposite to the phloem parts, but most common occurrence is
between the phloem and xylem. Root shortening (removing the root apical tips)
launches the branching. In practice, this is applied in pricking off, replanting small
plants or cutting of seedling roots, used to support rich growth of the root system.
As well as the above ground system, the root system grows for the whole lifetime
without limitations. Roots grow in length (primary growth) and in diameter
(secondary growth). In case of an adult plant the size of the root system remains
more or less unchanged. The ratio of dying off old roots equals the ratio of formation
of the new ones; the turnover is balanced. It is similar to what happens in the above
ground structures – treetop volume and the total area of foliage remains unchanged.
The lifetime is influenced by factors such as type of roots (anchor – long life,
elongated– long life, fine – short life), wood species and other conditions, both
endogenous and exogenous. Types of root systems of trees as widely described in
literary sources are generalised and regard individual species that live in the optimal
soil conditions. These root system types include superficial (plate) root system, tap
root system, heart-shaped root system. Yet, the variability of the root systems is
much higher than the changeability of treetops which corresponds to the fact that the
soil conditions are more heterogeneous and the shape or root systems change
during ontogenesis. For instance, a superficial root system may be observed in pine
(Pinus sylvestris) or beech (Fagus sylvatica) if they occur in swampy soil. Moreover,
it is frequent that the roots develop asymmetrically. Reaching of roots into the depth
of up to 2 meters may be even found in compacted and unsuitable belowground

32
conditions in case of young trees. Adult and old trees need to decrease
consumption of energy so they limit the diameter growth, lower treetops and
gradually move the root structure up into the higher surface horizons.
Root biomass makes up of about a third to a half of the biomass of the whole tree,
depending on site. The poorer the soil is the higher the root biomass share on the
whole tree biomass is. The division of the root biomass into various depths is really
interesting. E.g. Quercus family concentrates 90 % of the total root mass up to the
depth of 1 metre and 70% up to 40 cm. In case of conifers, the 30 cm surface level
concentrates 80–90 % of the root biomass. Generally, the surface horizon of about
40 cm in forest sites is considered a zone of intense rootage. However, in urban
conditions, this depth needs to be greatly regulated, even due to the intense soil
compaction.

4.2 Root functions

The root as an integral part of the plant organism holds the following functions:
anchoring the plant in the ground or adhering to the ground (mechanical function),
occupying new area by the root growing, uptake of water and mineral sources,
direct influence on the rhisosphere (material flow and direct control of conditions of
the symbionts development), disintegration of the rock, soil protection from
erosion. It also aerates the soil and enriches it with organic materials (due to root
turnover, i.e. dying and replacing with new ones).
Other essential functions involve the conductive function by transporting water
enriched with diluted mineral substances and gases, the storage function (starch,
mineral nutrients, water), the function of specified metabolism (esp. synthesis and
excretion) and information function (hormonal activity – root as a growth and
accumulation sing affects the production of photosynthates); protective function and
reproductive function (root stools) and the competitive function in view to other
organisms (root grafts, alelopathy etc.); it even has parasitical functions (haustoria).
If some of the functions prevail, the root inner structure as well as the general shape
may metamorphose.

4.3 Root zone activity (primary growth)

4.3.1 Meristem zone of the root primary growth

Roots grow in length on the root tips due to the activity of the initials of the apical
meristem. The initials of the rest centre demonstrate low dividing and metabolic
activity., New cells of the root cap, responsible for covering and protecting the root
tip, are divided on the distal surface of the rest centre. At the beginning, the root cap
cells are intensely divided, then they demonstrate a very intense metabolism and
gradually reach the edge of the root cap to peel off. Cells in the central part of the
root cap contain cascade amyloplasts. The change in their position tells us about
the direction of gravity. The cap excretes polysaccharide mucus, which makes it
easier for root to penetrate into the soil, protects the meristem from an injury, drought
or from separation from soil particles. It also controls the uptake of ions into the root
and supports the development of the microbial flora and mycorhizal fungi.
Cells of the resting centre divide their own initials of the apical meristem on the
proximal surface. By dividing (periclines and anticlines) the number of layers of root
cells is increased as well as the number of individual layers cells. This zone cells only

33
have primary walls, they are interconnected with plasmoderms and have juvenile
development features. A weakly developed membrane structure does not allow for
high speed of respiration, any differentiated xylem is absent. There are two main
functions of these cells: firstly, cell dividing as a precondition of the elongation
growth of the root, and, secondly, secretion of mucus enabling the penetration of
root into the soil.

4.3.2 The elongation zone

Cells divided from initials (derivates) undergo division as well as primary growth
in the elongation zone. The higher the number of cycles that cells complete, the
shorter they become in the end, and vice versa. Yet, not all parts of the elongation
zone divide at the same rate. This e.g. leads to developing many more epidermis
cells than vascular elements, related to the length unit of root. The heterogeneity in
cell division rate and derivates growth determines further histological and functional
root heterogeneity in the radial direction.
Water flow into vacuoles, containing glucose and fructose, is intensified, vacuoles
grow in size and fuse into a central vacuole. During the expansion, this vacuole
develops a pressure from the inside of cells on the cell wall. Their structures get
reorganized – polysaccharides split, new chains of cellulose as well as other
substances of cell walls are synthesised, such as hemicellulose, pectines, and even
some proteins. Not only the uptake of water is increased in the elongation root zone,
but also the uptake of nutrients – a part of whose gets accumulated and another part
gets utilized.
Cell elongation develops pressure in a distal direction on meristem as far as the
root cap, and in a proximal direction on differentiation zone cells with solid tissues. It
is this pressure on the root cap that allows root tip to penetrate into soil particles.
At first, roots grow through soil macropores (diameter over 0.1 mm); in case of a
risen soil resistance (compacted, sandy soils with the majority of pores below
0.001 mm in diameter) the secretion activity of the root cap grows substantially.
The period of root elongation depends on concrete species and it mainly occurs in
early spring when the soil has gone over the frost, and in late autumn. Daily growth of
roots varies greatly – it can be as little as 1 mm in Acer saccharum to as much as 56
mm in Robinia pseudoacacia. It is clear that a substantial role is played by the
physiological age of a tree. It is obvious that in young, newly planted trees the growth
is much faster than in the ones with well developed root system.

4.3.3 Differentiation zone

A process that has begun in the meristem is finalized in the differentiation zone. The
outcome of this process is shape and functional differentiation of cells and tissues.
On the root surface, there is a tissue called rhizodermis, which is a single cell layer
with no cuticle or stomata. The cells are rather thick from the exterior tangential
plane – there is a mucus layer on the surface, and a cellulose layer from the inside.
The mucus is not a secretion output as in the case of calyptra, but it is a part of the
wall. It contains a lot of negative ions and is enriched with microbial substances that
are important in an exchange of cations between soil colloids and the root.
Rhizodermis has a special capability of enlarging the surface by the creation of
root rhizins; this takes place roughly at the same distance from the apex as the first
functional differentiated elements of xylem. Rhizin is responsible for a uptake of

34
nutrients and for a mechanical support. Such a rhizin activity is not as usual and
intense in trees as in herbs, the rhizins are shorter and sparser, but they have longer
lifespam than in herbs. It entirely lacks in unfavourable soil conditions, esp. weak
aeration, or if a root is occupied with mycorhizal fungi which fully compensate the
rhizin function. Older rhizodermal cells and rhizins suberinize, peel off and are usually
replaced with secondary cork.
Under the rhizodermis, there is primary cork – much thicker in root compare to the
central cylinder – except for e.g. Vaccinium which is single-layer. Primary cork of
trees is made of mesoderm, often containing wide intracellular spaces (aerenchym)
developed mainly in wetty soil conditions and a lack of oxygen. Exodermis is not
formed in many cases at all. Besides the aeration function, the primary cork
parenchyma also has the following functions: storage function (starch, proteins,
lipids), excretion function (accumulation of tannin, mucus, ether oil and crystals),
protective function (accumulation of toxic substances and detoxication of these in
vacuoles), synthetic and transport functions. All these functions are partly held by
mycorhizal fungi that live and develop in mesoderm. A lot of substances like
water, nutrients and gases, e.g. carbon dioxide or oxygen are transported from the
soil through these cells through the apoplast – by diffusion through permeable cell
walls and intracellular areas and by the symplast – through life cell bodies.
Endodermis cell walls are suberinized and lignified in the radial and transversal
directions (Casparian strips). These strips prevent from further transport through
the apoplast so most transported matters have to cross the barrier of
semipermeable membranes (plasmalems or tonoplast) and enter the symplast. The
local suberination and lignification of cell walls is only absent with permeable
transfer cells located opposite xylem groups of the central cylinder. Roots that
demonstrate secondary diameter growth retain rhizodermis and primary cork only
temporarily, they can be found in fine roots and the youngest parts.
The term central cylinder stands for a column of conductive tissues system inside
the root, on its periphery bordered with pericycle. In more primitive gymnosperms
the pericycle has multiple layers as a whole, or multiple layers may only be set locally
opposite xylem and phloem groups. The pericycle cells retain the capability to
divide – they take part in regeneration processes, they are a formation place of
lateral roots and part of cambium, sometimes even phellogen. Their cell walls are
penetrated with plasmoderms securing a contact between cells from endodermis and
parenchyma in the central cylinder. The conductive system in root is represented by
a complex radial vascular bundle whose individual parts of protoxylem and
protophloem are arranged in rays. Phloem and xylem sections alternate. Based on
the number of phloem and xylem groups, bundles are categorized into monarch,
diarch and triarch (gymnospermous and dicotyledonous) and polyarch
(monocotyledonous; palm trees, Monstera). Nonetheless, this rate may vary even in
just one individual, such as the case of Picea albies with diarch, triarch bundles and
monarch vascular radial bundle in its short roots.
Nearest to the root tip (0.1 – 0.4 mm) there are differentiated elements of phloem that
take photosynthates and required K+ into the system. Further away from the root tip,
in an area where a root starts to operate as an organ absorbing water and nutrients,
there is xylem. In Abies fast growing roots it is located 7 mm from the apex, in Acer in
inadequate conditions it is only 0.2 mm. At first, xylem and phloem groups are
arranged radially and their number is unstable. Xylem is differentiated in centripetal
direction (exarch): protoxylem elements are closer to the perimeter of the central
cylinder, and towards the centre there are metaxylem elements and there is

35
usually no pith. Conductive pathways of wood (especially vessels and tracheids)
act as dead elements. Intense synthesis of cell wall substances (cellulose,
hemicellulose and lignin) is active in their protoplasts during the elongation and
differentiation. After the construction of these is finished, protoplasts disintegrate.
Substances released at the programmed death of these cells become part of
the whole transpiration flow and enrich nutrition of the above ground system.
Roots get branched behind the differentiation zone: root primordia are initiated
endogenously in pericycle. Because of this, they do not have an effect on the
structure of the conductive system of the parent axis, as they do in the case of stem.
When growing they tear the outer tissues of the root till the soil is reached. Even
roots with suberinized surfaces demonstrate a relatively high uptake of matter at
places of root branching. Root branching multiplies the number of root tips, allows for
a higher density of rootage and more intense utilization of a suitable soil area.
Apical, distal parts of roots are the most active parts in terms of synthetic
activity, impact on rhizosphere, water and nutrients enriching from the soil.

4.4 Secondary growth and root functions

Roots of a tree grow into the depth by means of two types of lateral meristems. We
can think of them as single layer shells of concentric cylinders. In the case of
gymnosperms and dicotyledons, cambium is initiated in the central cylinder
(proximally from the apex).
The first primary parenchyma cells that undergo the remeristemation are located
at the inner side of the primary phloem, then on the boundary between phloem
and xylem groups and after that in the pericycle, located at outer sides of
primary xylem groups. Cross sections show that the layer of cambial cells could be
oval (diarch roots), triangular (triarch roots) and so on in their early stages. Ensuing
from the fact that cambium cells gradually divide mother cells of secondary xylem
centripetally, mother cells of secondary phloem centrifugally and cells of secondary
phloem-xylem rays between bundles in both directions, cambium generally
becomes of a circular shape on the cross section (Fig. 16).

Fig. 16 Cross-section of Norway spruce fine roots. Vascular bundle was radial
(diarch) at the early stage of fine root development. Primary xylem (1 – protoxylem
and 2 – metaxylem); 3 – secondary xylem – first grow ring; 4 – resin ducts; 5 –
secondary xylem – second grow ring; 6 – secondary phloem; 7 – pericycle; 8 –
primary cork

36
Cambium in roots is active at different time in a year than cambium in the above
ground systems. Cambium may demonstrate different activity even among roots
in one individual. The same may happen even within one root – in one root part, it
may be more active than in others. This is mainly due to mineral components present
in the soil. Root diameter increase in heavy and sandy/rocky soils is irregular;
most roots incline to develop eccentric structures, which further limit their
conductivity. Annual rings in roots are thinner than in stems, they are hardly
visible, and a typical clear layer of early vessels is missing in the ring porose ones.
The proportion of individual elements of xylem in roots rather differs from the ones in
stems and branches. The secondary xylem in roots can be characterised by
intense development of conductive pathways (vessels and tracheids) and by a
higher content of parenchyma. This allows for easier transport and storage and it
supports the active participation of roots in the processes of material exchange. The
number of xylem fibres is reduced, except for the ones in anchor roots and the
ones with high mechanical requirements. In general, cells in roots are bigger than
in stems and trunks; in tangential and radial direction as well as in length.
Phloem conductive pathways remain active or dormant in winter, but they usually
degenerate as early as the end of their first growing season. In spring, new phloem
elements differentiate from derivates of cambial initials, having overcome winter in an
imperfect state.
Along with a diameter increase of secondary roots, another secondary meristem,
called phellogen, is formed. It is made by the remeristemation of pericycle cells.
Secondary meristems and their derivates could react to changes in the root
environment. For instance, if aeration is reduced, abnormal xylem and phloem with
an increased content of parenchyma are formed, transport of carbonhydrates is
inhibited, absorption of macrobiotic nutrients is reduced, root metabolism is limited.
For example, hypertrophied lenticels are formed and the generation of
adventitious roots is supported in some species (Salix alba).
Even though the secondary root structure may considerably limit the intensity of
nutrient uptake from the soil, the total uptake is not insignificant compared to the
activity of root tips. Although secondary roots have suberinized surfaces this surfaces
have fewer, thinner and more permeable layers than in the above ground system.
Moreover, they are connected with the conductive system through phloem-xylem
rays. Other locations with an intense exchange with the soil include disturbed areas
of suberinized surfaces near lateral roots initiations. Regarding that the total surface
of suberinized roots is very large, the uptake of soil solutions through this structure
could not be omitted. If roots are uncovered and exposed to air for a long time, their
bark or rhytidome resemble bark of a trunk both in the visual and functional aspects.
Such roots lose the uptake ability and they mainly function is mechanical support and
conductivity.

4.5 Mycorrhiza

The term mycorrhiza (from Greek mycés – fungi, rhizó – root) refers firstly to root
symbiosis of higher plants with soil microscopic fungi, secondly to organs formed
during the symbiosis of a host plant root and an endophytic fungus. The latter is
mainly used in ectomycorrhizae, in which the association of a root with a mycorrhizal
fungi substantially affects morphological and anatomical root structure.

37
Mycorrhiza are the most widespread symbioses that roots of plants create.
Foundations of plant roots fossils indicate that the predecessors of present fungi
forming vesicular-arbuscular mycorrhiza lived in the symbiosis with plant roots as
early as their expansion from water to land environment, which dates back to approx.
350 – 460 million years (from the Ordovician to the Carboniferous periods). Other
mycorrhizal types are younger and more derived.
One of typical features of mycorrhiza is that their development in roots of host
plants is only limited to the primary cork of roots. Mycorrhizal fungi never appear
in the central cylinder. Another distinctive feature of all types of mycorrhiza is a
formation of a network of mycelium in soil, not in roots. This extramatrical
mycelium, functioning mainly as a satellite of a plant root system, substantially
enlarges soil volume as a source of nutrients, esp. phosphate ions, to the plant.
Based on the latest monograph dedicated to mycorrhizal symbiosis, the following
basic types of mycorrhizae are distinguished: orchideoid, ericoid, arbutoid and
monotropoid. The older categorization into endomycorrhizal and ectomycorrhizal
symbiosis related to the capability of a mycorrhizal fungi to colonise cells of
primary cork. Only the endomycorrhizal are able to do this. The development of
an endophytic fungus in ectomycorrhizae is limited to intracellular spaces in
primary cork. In this aspect, ectendomycorrhiza, arbutoid and monotropoid
mycorrhizae are a transitional type –fungus colonises primary cork cells, but these
mycorrhizae have some common features with ectomycorrhiza.
The following text desrcibes only the types of vesicular-arbuscular mycorrhizae,
ectomycorrhizae and ectendomycorrhizae as these are the ones that play the most
important part in woody species.

4.5.1 Vesicular-arbuscular mycorrhiza (VAM) = arbuscular mycorrhiza

(AM)

VAM includes a really wide spectrum of species of host plants – it is estimated that
about 80% of all plant species form this kind of mycorrhiza. In this view, it is the most
widespread symbiosis in the world. It can be found in plant species through all
climatic zones and environmental conditions, no matter if they occur with trees,
bushes or herbs. Typical woody species that form VAM are from Cupressaceae and
Taxaceae families; VAM also occurs in seedlings of Pinaceae family, though this
family usually forms ectomycorrzhizae.
No species-specific VAM symbiosis has been documented yet, so, it is generally
presumed that there is no such specialization. In a different manner, there has been
an evidence of a certain kind of environmental differentiation. It lies in the preference
of certain fungi species by different plant species in the same environmental
conditions.
Typical structures of this type of mycorrhiza are arbuscules, alternatively vesicles,
whose presence makes the background for the most widely accepted definition.
Arbuscules are formed inside cells of a root primary cork. They could be
characterized by their multiple fork-like branching and finalized growth. Arbuscule
cell wall is thinned or absent. The plasmatic membrane of a fungus is never in a
direct contact with cytoplasm of a cork cell. Arbuscules are considered to be the
centre of symbiotic transmission of phosphates and carbohydrates between a fungus
and a plant. Vesicles are located either inside root primary cork cells or in
intracellular spaces. They have a storage function, containing mainly lipids.

38
4.5.2 Ectomycorrhiza (EM)

In comparison to VAM host plants, the number of EM host plants is very small – it
only makes up about 3% of species of all plants. Yet, this is an especially significant
type of symbiosis in our tree species as it appears within families Pinaceae
(especially Pinus and Picea) and Fagaceae (Fagus and Quercus). Another important
families with EM are Myrtaceae in subtropical and temperate climates in the southern
hemisphere and Dipterocarpaceae in the monsoon forests in the South East Asia.
Even other species of other families form EM (e.g. Betulaceae). Some species can
either form VAM or EM, depending on soil conditions of the site. Such families
include Eucalyptus, Salix or Populus – EM is formed in conditions of very humid
soils.
Compared to VAM, when the creation of mycorrhizal association does not affect
morphology of an infected root, the formation of EM symbiosis leads to
substantial changes in the structure of root tips (only these are capable of EM
formation). Their apical meristem finalizes its growth so root is shorter and thicker.
The extent of the shortening and thickening processes depends on the development
stage of a root tip at the moment of its contact with a mycorrhizal fungus. Another
typical feature is forky root branching. An ectomycorrhizal root has several fold
longer lifetime (approx. 3–4 months) than a non-mycorrhizal root tips.
The following structural components are typical for ectomycorrhiza:
1) Fungi mantle – it covers root surface. It may only be weakly developed or
absent. Typically the biomass of a fungus mantle makes up of 20 to 40 % of the
total biomass of a short EM root.
2) Hartig net – a labyrinth of hyphen growing from fungus mantle into the
rhizodermis and intracellular spaces of root primary cork. It is typical for these
hyphen to be branching intensely. Spreading of hyphen may be only limited to
rhizodermis, (typical for angiosperms, or they may grow in intracellular spaces of
more layers of the primary cork, they sometimes might grow through all layers of
the primary cork (typical for gymnosperms). Hartig net is a place of the most
intense material exchange between associated symbiotic organisms.
3) Extramatrical mycelium – serves to absorb nutrients. It may be aggregated
into rhizomorphs, i.e. thick strands of hyphen, that are clearly visible.
Rhizomorphs serve to transport material on long distances.
Older studies regarded fungus mantle as a centre of uptake of nutrients from the soil.
Nonetheless, at present, the extramatrical mycelium is considered to take an
essential part in the absorption of mineral substances as in the case of other types of
mycorrhizae. The Hartig net is a place where minerals absorbed from the soil are
exchanged for carbohydrates assimilated by a plant in the photosynthesis.
Ectomycorrhiza is mainly formed in the soils rich in organic substances,
especially surface horizons of forest soils. A substantial portion of phosphates
and other nitrogens in these horizons may exist as organic compounds. EM fungi are
able to disintegrate these organic compounds by means of hydrolytic enzymes, and
perhaps even due to associated bacteria. Acceptable mineral forms of nutrients for
plant are formed.
4.5.3 Ectendomycorrhiza
Although ectendomycorrhizae are quite widespread, we do not have much reliable
information on them. They could be formed by woods that commonly form
symbiosis with EM, especially in the early ontogenetic stages (described e.g. in

39
Pinus, Larix, Picea). Some fungus taxons with active ectendomycorrhiza are related
to EM fungi, as well, since these may form a typical EM, depending on a host plant.
All aspects of development and formation of typical structures are comparable to EM,
except for the intercellular penetration of primary cork cells of a host root. Both
fungus mantle and Hartig net are created. And, at the same time, hyphen grow
through primary cork cells where they are intensely branched. This symbiotic
association is very stable, and, similarly as in EM, roots do not demonstrate any
traces of senescence.

4.6 Metamorphoses of root

In the course of evolution, root functions have changed considerably in many

instances. This concerns all types of roots – main, lateral or adventitious. Along with
the change in function, the shape and inner structure have changed, too. Root
metamorphoses may be so enormous that the original character of this belowground
structure has gone completely away.
Many adventitious roots of species located in loose or muddy soils fulfil mechanic
functions. It is as if plants were standing on stilts, that is why these roots are called
stilt roots. The same function is also taken by so called prop roots that support giant
trees of many tropical species, and buttress roots, developed in some species in
tropical rainforests. Many lianas, e.g. local Ivy (Hedera) and numerous epiphytical
species are fixed by adventitious roots. The fixation function is also made by
adventitious roots, formed on stems of creeping species, e.g. Cotoneaster, Salix,
Pinus mugo and the like. A plant uses them to be tied to the soil, and at the same
time to take in nutrients. They can even be separated by means of the vegetative
reproduction an form a polycormons. Some spines on palm trunks originate from a
root.
Aerial roots commonly appear with epiphytes. They grow freely in the air and their
dead cells of multilayer rhizodermis absorb water from the rainfall. Such roots may
also have the assimilation function. Species that live in wetland areas with a lack of
air have breathing roots, pneumatophores, that mediate an exchange of gases.
They grow right up in the air, their tips stick out of the soil or water (Taxodium
distichum).
Parasite roots (haustoria) are special case of metamorphoses. Some lateral roots
develop as tiny tubers. Haustoria, meaning organs that draw nutrients from a host
plant, grow from the tubers and penetrate as far as vascular bundles of a host
(Viscum, Loranthus).

4.7 Limiting factors of root system growth and functions

Firstly, these are the factors that reduce the positive carbon balance in the above
ground system (i.e. balance between the photosynthesis and respiration) or
negatively impact the translocation of photosynthates and growth hormones in a
different manner: radiation stress, temperature stress, water stress, intoxication,
injury stress (by an intentional cut or by the wind, snow, frost), oxidation stress (by
increased entry of oxide substances such as O3, SO2, NOx, PAN etc.), damage to
conductive pathways and tissues (by frost and bark scarlatina of unshaded
trunks), by fungi, viruses, insects and a range of other negative factors including
the ones that are connected with various defoliations.

40
Other important factors include soil factors, especially the soil type, into which roots
penetrate and regenerate. Further, it is a high content of skeleton (rocks, stones,
remainders of brickwork etc.) and the mechanical impedance of soil that limits the
root penetration. The soil in urban areas is commonly compacted (by walking,
driving of heavy machines) and flooded (as water from rains cannot easily get
absorbed). If soil aeration is limited (hypoxia), this leads to a higher production of
potentially toxic materials, to the modification of shoot growth, inhibition of formation
and growth of leaves. When this affecting is long-term, the diameter growth of adult
trees slows down, and the phloem and xylem anatomy gets transformed. In such
cases, the absorption of minerals is limited due to a smaller extent of the root
system and a lack of energy. This is because of the fact that in the anaerobic
respiration of an organic substrate, less energy is released than is required to cover
the needs of complex root functions. Not only the soil drought, but also hypoxia may
stop the uptake of nutrients and it may even happen that ions are released from the
roots out into the soil.
The soil drought results into a worsened uptake of water by the roots, transpiration
and assimilation limitation (i.e. worse cooling of leaves, supply of leaves with
nutrients and roots with photosynthates). Moreover, it results into the efflux of
nutrients in fine roots and into the limited contact of roots with the rhizosphere, and
may end in death of roots. The soil drought together with unfavourable atmospheric
conditions such as low relative humidity, high temperature or wind; generally the
ones that lead to an increased evaporation, might bring about a damage to the
conductive pathways in xylem by high negative pressure. This may cause breaking
of water columns and a consequent suction of air or cells (from the neighbouring
parenchyma) into the vessels.
Fine roots mortality may also rise due to a high content of salts in the soil, due to
an imbalanced uptake between individual nutrients, low recoverability of nutrients
and an absence of mycorrhizae, winter starvation, summer temperature (thermal
convection of asphalt surfaces, thermal radiation of buildings etc.).
One of ways of sustaining trees in towns is to secure them conditions needed for
their well being. This means to make a range of tasks, namely to secure long-term
good quality and volume of the belowground and air space, and to regularly secure
their requirements for water regime (irrigation, evaporation), temperature regime
(planting in groups) and light regime (i.e. to respect demands on density and quality
of radiation, length of day and growing season, sum of radiation in a growing season
and in a calendar year) and to regularly manage natural soil processes (i.e. mainly
the recoverability of nutrients and the presence of suitable soil flora and fauna).
In confined urban spaces such as pedestrian zones or stations of public transport
etc., trees are often placed into special containers on the ground level covered
with decorative lattices, serving as prevention from the soil compaction by walking.
These lattices should be made of modules dismountable in the radial direction (in
case of a round container) as well as the tangential direction (links between partial
circles). Such a treatment is especially required by the base part of a trunk, since the
volume changes in the diameter growth are really great at this place. The same is
true for volume changes in the root system due to the elongation growth. The base
part of a trunk may quite soon reach considerable volume changes at the places of
connections of roots with a trunk, which are of a high importance for the mechanical
support of a tree. The ground level containers have the following disadvantages:
easy entry of salt water from melting snow and ice removed from roads. The salt

41
solution in winter has a much lower temperature than the soil, which means that it
may promptly and repetitively contribute to freezing out of fine roots.
In the urban zones, trees are also held in above-ground containers placed on the
surface of streets. They have these great advantages – the soil is protected from dirty
salt water and from being walked on. On the other hand, the soil is not protected
from the frost at the beginning and end of the growing season, or from overheating
and drying in summer.

5 The stem

The stem (caulom) is a vegetative, usually above ground structure with nodes. It
generates, differentiates and holds lateral branches, leaves, flowers and fruits.
The stem mediates their connections with the roots by distribution of water,
nutrients, photosynthates, and material, electric and stress information signals etc. It
participates in photosynthesis, defence and protection, its directed growth
occupies the optimal space and it is a storage of water and starch for
overcoming bad periods. The elongation growth, generation of leaves and lateral
branches is made by the apex meristems. The connection of these new modules to
the roots is secured by the cambium. A significant role of a stem is also to maintain
spare suppressed buds that can renew growth after a heavy stress, e.g. after an
injury by wind, lightning, browsing, frost, drought, starvation etc. Some other
important roles include vegetative reproduction or holding a plant on the spot.
Internodes of a stem are sections between two nodes, which are a base of leaves
and buds. The length of internodes corresponds to the biological type of a plant
(longest in lianas, shortest in gymnosperms – on shortened shoots holding
bundles of needles – brachyblasts), to the age and position in a tree crown (e.g.
quickly growing seedlings after a deep reduction of a tree crown compared to edge
parts of crowns of old trees), environmental condition (shaded stems demonstrate a
lower length of internodes than stems in the direct sunlight). A tree stem with leaves
is called a shoot.
An unbranched part connecting the root and the tree crown is known as a trunk,
stems within the crown are called branches. Trunks usually have the radial
symmetry, the most frequent shape is cylindrical, but may also be muscular, flattened
etc. At the bottom, it is extended by roots. By the direction of growth and self-bearing
capacity, stems are categorized into different groups: direct (heliotropic – grow up
vertically, plagiotropic – grow horizontally, bent when older and grow down),
overhanging (pendular), ascending, lying, creeping (cotoneasters), supporting
(blackberries) etc. Trunks are among the greatest organs in the plant kingdom. Palm
Calamus (leaf-climbers) might reach as high as 300 metres. The tallest trees reach
up to 150 metres. These are Australian eucalyptus which are even taller that sequoia
(Sequoiadendron giganteum), reaching 120 m. The greatest circumstance has
Mexican Cypress (Taxodium mexicanum). With the diameter of 31 m it wins over
North American sequoias. Nonetheless, sequoias still hold the world record as the
largest organisms by volume.
The stem structure is made of covering tissues, primary tissues, conductive
tissues differentiated from the apex meristems. In this case, the epidermis on the
surface is a single layer (sometimes with well developed stomata), it has commonly a
cuticle and trichomes. In a multi-layer epidermis tissue, layers below the
epidermis are called the hypodermis. The primary cork lies under the epidermis

42
and consists of two to three layers – upper layer of primary cork has a mechanical
function. It contains collenchyma, commonly with chloroplasts (in young stems it
quickly loses its tension (turgor) when water is absent). The medium layer is usually
parenchyma with the assimilation function. The lowest layer is made of living cells of
endodermis or its cells of another kind, such as starch stripes, but it could be also
absent. The primary cork has mechanical, storage, assimilation and protective
functions..
Closer to the centre, there is a central cylinder. On its periphery, there is a
pericycle (latent, secret meristem where adventitious roots and buds may originate),
below, there is a system of conductive tissues, i.e. vascular bundles. There are two
types of vascular bundles in stem of gymnosperms, dicots and monocots (Fig.
17). The first lead from the stem to leaves, the other run through the stem
individually. In monocots, all vascular bundles are leaf traces. In case of conifers
and angiosperms, the vascular bundles are collateral, arranged in a circle. On the
cross-section they create a circle surrounded by the primary cork at the outer side,
and by pith at the inner side. Pith is connected with primary cork by stripes of
parenchyma tissues, pith rays.

Fig. 17 Vascular bundles arrangement in gymnosperm and dicots (left) and monocots
(right).
Stem pith fills the centre. It has thin walled cells, most commonly freely connected,
cell wall is sometimes weakly lignified. Intercellular space in pith are filled with air or
water. Pith of some trees quickly breaks (Juglans, Vitis vinifera) and new cavities are
formed which are interrupted by partitions. While in some species, pith is sustained
for a long time and functions as an assimilation tissue (ash tree), in others it dies after
the first year, e.g. elderberry bush.

5.1 Secondary growth of stem

Secondary growing stems gradually develop a secondary structure depending on

the formation and activity of secondary meristems (Fig. 18). These include cambium,
which replaces the systems of conductive tissues divided into phloem and xylem
sections of vascular bundles, and the systems of primary tissues with phloem-xylem
rays, and resin canals in gymnosperms such as spruce or pine. We distinguish
between two types of cambial initials in woody plants. Cambial initials are named
fascicular cambium (form xylem and phloem. The second type, interfascicular

43
cambium (form pith rays), is usually formed at the end of the first year. Fascicular
and interfascicular cambium together make a continuous cylinder on the cross
section. Another secondary meristem is phellogen, which generates secondary
covering tissues, i.e. secondary bark.

Fig. 18 Cross-section of one-year old Norway spruce shoot. 1 – pith; 2– primary

xylem; 3 – secondary xylem; 4 – secondary phloem; 5 – pericycle with resin ducts; 6
– primary cork; 7 – resin duct

5.2 Development of annual rings

Cambium activity depends on the activity of meristem cells of the stem apex, lateral
buds, signals from leaf, roots and suppressed buds. It begins in spring by first
dividing and differentiating of phloem and xylem mother cells that were formed in
the previous autumn and survived winter. Cambial initials take water into vacuoles,
their thick cytoplasm is pushed by the growing vacuoles to the wall which is
extending and becoming thinner in the radial direction. This action begins below
buds due to the phytohormonal factors (auxins, gibberellins, cytokinins with co-
working of carbohydrates).
In the case of the diffuse porous woody plants, this activity directs from stem tips
towards the trunk base. If ring porous woody plants are considered, this process
begins later, but is faster, and as early as the first buds begin growing, the first
vessels are functional. Regarding conifers, the impulse comes from old as well as
new needles. Dividing of cambial initials begins a couple of weeks later, the division
spreads all over the trunk and the process ends in the root system. The same
process applies for closing of cambial activity in autumn, and this is why roots grow
even in autumn.
The first cells produced by cambium in spring are xylem mother cells, phloem
cells follow. In this period xylem cells grow really quickly and their cells have larger
lumen and thinner walls. Such wood is important for its conductive function. In
summer, the situation is reverse. Cambium activity slows down and latewood cells
are thick-walled; they have mechanical and defence functions. So this is how early

44
and late wood originate – its annual growth is known as an annual ring (Fig. 19).
However, there are cases when more than one annual ring may be formed in a year.
Such false rings come from a defoliation of a tree (e.g. by insects), interruption of
the growth activity (e.g. by drought) and due to other unfavourable factors. The
activity of phloem formation is weak in spring, but it sharply rises in the course of
the growing season and continues till autumn. In spring, the cambial zone is thick
(cells of initials with mother cells of conductive and primary tissues), in autumn, it is
thin, on the contrary. Cambium activity in young trees starts earlier in spring and
finishes later in autumn than in the case of old or weak trees.

Fig. 19 Cross-section of 13-years old Norway spruce branch. Annual ring is formed
by earlywood (conductive function) and latewood (mechanical and protection
function)

5.3 Wood of coniferous species

It is an evolutionary older type. It can be characterised by a simple structure, typical

alternation of early and late tracheids and parenchyma cells (Fig. 20). These
elements build the structure of annual rings. The main constituent of wood is
tracheid, that makes up about 90% of volume of coniferous wood. Conductive
function of early tracheids reflects their shape – thin walled, prosenchymatic
cells, length 2–6 mm and diameter 0.04 mm (wall thickness 2–3 μm). Water flows
through border pits in the cell wall, which is why coniferous wood is quite
resistant to water flow. Water is well supplied due to an increased rate of early
tracheids – trees in good years form more early wood than late wood. The wider the
annual ring is the more early tracheids are produced. Yet, tracheids fulfil the
conductive function longer than vessels of deciduous species, and a larger part of
wood is active in water transport (more annual rings). Late tracheids are thick-
walled, sclerenchyma cells. They are roughly 10 % longer than early tracheids,
and cell wall is 3–7 μm wide. Another cell types are parenchyma cells. These are
thin-walled, isodiametric cells that play a role in the formation of resin canals and pith
rays. They fulfil secretion functions, secure radial transport of water and
substances, they fulfil also mechanical functions.

45
Fig. 20 Coniferous wood structure (Picea abies)

5.4 Wood of deciduous species

The wood of deciduous plants is phylogenetically younger than the wood of

conifers. This is reflected in a higher complexity of the structure, a higher
assortment of cell types, which are more specialized on certain functions. Deciduous
wood is made of vessels, tracheids (fibre, vasicentric, tracheidal), libriform fibres
and parenchyma cells (Fig. 21). Vessels are specialized conductive elements
made of vessel elements. They may reach considerable lengths (5–18 m for oak,
0.8–2 m for birch). The most common length is 10 mm. Taking into account their
considerably thin cell walls, vessels do not have a high mechanical stability.
Therefore, they are reinforced with spiral or round thickening. Depending on
species, the rate of vessels in the overall wood volume varies from 4–8 % (maple) to
22–48% (Populus, Fagus, Fraxinus). There are three types of deciduous
tracheids: tracheidal, vasicentric and fibrous. The first ones mainly fulfil a conductive
function and do not occur with the majority of deciduous trees. Vasicentric tracheids
also have a conductive function, but only appear in company with vessels, such as
oaks. Fibrous tracheids are cells with a mechanical function, but they can also fulfil
conductive and storage functions. Libriform fibres are mechanical elements of wood
of deciduous trees. Their percentage representation ranges from 36% (Fagus, Tilia)
to 78% (Acer). These are thick walled, short, cylindrical or spindle cells. Their shape
and size make them really stable cells that can secure the wood solidity. Parenchyma
cells in deciduous cells make up about 8 – 35% of wood volume. They are tiny cells,
mostly isodiametric, sometimes of a roll shape.

46
Fig. 21 Structure of deciduous ring porous wood (Salix fluviatilis)

5.5 Stem morphology

Trees have a distinguished main stem, trunk. It does not branch and usually bears
lateral branches and leaves. Skeleton branches (branches of the first class) bear
macroblasts (branches of the second and higher classes). Shoots with shortened
internodes are called brachyblasts. In some species, they are formed regularly
(Ginkgo, Larix), but in most species only as a modification which allows for the
annual generation of new leaves, flowers and fruits without extreme growing of a
crown (Tilia, Fagus, Malus). All branches make a crown. It gives trees their typical
appearance – habitus, depending on means of branching, number of branches,
direction of branches, foliation etc.
Monocotyledons have enclosed vascular bundles, they are scattered and all of
them pass from stem to leaves; there is no boundary between the primary bark and
the central cylinder, there is no pith or rhytidome. Secondary thickening is rare
(does not exist in palm, bamboo). It is carried out through cambial initials centrifuginal
dividing of primary parenchyma tissue (gradually sclerifying) and individual vascular
bundles. The phellogen is formed on the outer side, dividing cork, with an occasional
occurrence of parenchyma and sclerenchyma cells (aloe, dracaena, yucca).

Branching of tree stems is holoblastic (apex tip does not split, it remains as a
whole complex). There are two types:
1. Monopodial – racemose. The main apex is the tallest, lateral branches are
shorter. It is thicker and its individual modules are derived from the original apex tip.
2. Sympodial branching. The main apex is the shortest and lateral branches
are taller. Sympodium is further classified into:
a. monochasium, formed in woody species with alternate leaves and
node buds. At the end of the growing season the apex growth slows down
and the tip bud either remains under-developed or does not survive winter.

47
 In late summer or the following spring, the nearest lateral bud gets into the
position of the terminal and gives rise to a new main shoot. By growth of
lateral branches of higher and higher classes, a branch elongates (Tilia,
Betula, Populus). Monochasium is also developed in wine (Vitis vinifera),
whose apex bud metamorphoses into a shoot. A lateral bud – based in a
leaf node overtakes its leading position.
b. dichasium is typical of woody species with an opposite position of
leaves and node buds. Even in this case the leading stem weakens as
time passes, or the terminal bud differentiates as a flower. Then it shoots
and there arise two almost equal stems of both opposite buds (Syringa,
Acer, Aesculus).

Monopodial woody plants rarely retain this exact type of branching in their lifetime;
they often pass to the other branching pattern. There are a lot of factors leading to
this change, like injury, parasites, inadequate functioning of the root system, drought
or age of an individual. Monopodium remains the main type of branching for a long
time for instance in spruce with a conical crown. Yet, its branches incline to
sympodium. Similarly, a crown in young pine trees is monopodial, but old pines
crowns are not monopodial, which reflects in bizarre shapes of crowns. The
sympodial branching, especially dichasium, forms rounded and oval tree crowns
or conical and conoid crowns. The shape of a crown is more plastical and can better
capture the sunlight and prevent from evaporation. On the other hand, if a dichasium
branching develops too soon, two main skeleton branches may develop in the crown,
whose phototropic growth (towards the sunlight) drives them apart, and this may end
in a huge break due to the opposite forces of two large heavy branches. The tree
crown, its size and shape are defined by the species and age of a tree, i.e.
differences in rate of growth of terminal and lateral branches and type or branching.
However, even sunlight (solitary crowns and individuals in related greenery),
unidirectional dry wind (flag-like crowns) and low soil humidity (umbrella crowns)
play a significant part in the final crown shape.

48
Fig. 22 Stem branching pattern and tree and shrub growth. Monopodial – the main
apex is the tallest, lateral branches are shorter. Sympodial branching – the main apex
is the shortest and lateral branches are taller. Acrotony – growth form of tree.
Basitony – growth form of shrubs.

The arrangement of leaves on the stem, phyllotaxis, is defined by plant species. In

the opposite pattern, leaves grow in pairs facing each other, turned of 90° from the
lower node (e.g. Acer, Aesculus). If more than two leaves grow from a node, this
pattern is called whorled (e.g. Catalpa). The position in more or less loose screws is
called alternate (or spiral). In this case, some of the leaves are positioned in vertical
lines (orthostichies). If we want to get from a given leaf to the nearest in the same
orthostichy, we need to go round the stem several times and pass leaves of other
orthostichies. The number of round and leaves defines the type of a leaf pattern.
Distichal (1/2, meaning two leaves in one round, the third leaf in the same line above
the first) is typical in shaded branches, such as in beech, elm, lime and monocots.
Dicots bear leaves in the pattern 2/5 (but also 3/8, 5/12 etc.). If the arrangement of
leaves is modified by their light reaction, i.e. change of spacial arrangement e.g. as a
result of a leaf mosaic, it is necessary to study the position of leaves in a bud to be
able to define the type of phyllotaxis. The shape of stem pith relates to phyllotaxis.

5.6 Bud

The bud is the foundation of a future structure (shoot, leaf, flower). It consists of
apex tip meristems, leaf primordia in the acropetal order, and axil buds. The oldest
leaves, sometimes metamorphed into scales, protect a bud.
Buds at the tip of stems (branches) are called terminal buds. This is the leading
apex bud of the main stem or trunk. Buds in the axils of leaves, i.e. in stem nodes,

49
are axillary, lateral buds. The vertical arrangement of these is called serial
(Lonicera), horizontal is called collateral (Prunus spinosa). Apex and lateral buds,
i.e. regular buds in trees flowering prior to foliating, may be subdivided into flower
buds bearing the base of a shortened stem and leaves modified into flower parts,
and leaf buds bearing the base of a shoot. Undifferentiated buds are mixed, giving
rise to shoots first and flowers later (Tilia).
The classification by the inner differentiation defines buds of the fir and poplar type.
Regarding the fir type buds, the whole shoot foundations were based a year prior to
spring when they begin growing in volume. Considering the poplar type, new shoots
develop according to climatic conditions in the same year of shooting and growing,
there is no significant role of the previous year.
The buds that are passive only for one resting period and start developing the
following spring, are called winter buds. The ones that remain passive are called
dormant buds. If these grow and branch, but still do not reach the surface of the
bark, are called suppressed buds. This type enables the reproduction and
regeneration of a wood plant when its integrity is violated, perhaps by the frost, fire,
weak functioning of conductive tissues, age etc. The same function is held by the
adventitious buds, formed beside the regular order on the internodes, nodes of a
stem or roots. These reserve buds are commonly called preventive buds.
The inner structure as well as the outer appearance could be considerably modified
or metamorphed from the above described models if a stem takes over other
functions, such as a defence function – prickles. Some species have prickles only in
young individuals, or root sprouts or low branches, while they are absent in a crown.
The climatic conditions play a part too, e.g. in blackthorn these structures only
develop on dry and warm sites, while in humid conditions of a greenhouse, they do
not develop at all. This is due to the fact that in this case thorns etc. are a protection
from herbivore animals. Assimilation (green) stems have e.g. Cytisus and stems of
succulents serve as reservoirs of water. Stems also secure vegetative
reproduction when branches are in contact with the soil they might give rise to
adventitious roots (Fagus, Picea aj.), which could later develop into individual plants.

Fig. 23 Leaves and axillary buds are formed in the first year. Terminal buds are
formed when shoot elongation growth slowdown. Conifers form axillary buds only at
the base of few needles.

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6 Leaf

Leaf (phylum) is a lateral structure generated by the stem, it has a pre-defined

position, limited growth and age. A leaf fulfils numerous functions: photosynthetic
(i.e. reception of solar radiation, CO2 and release of oxygen), transpiration,
conductive (i.e. evaporation, cooling, control of cell tension – turgor, supply of leaf
with nutrients and conduct of photosynthates away) and respiratory (i.e. respiration
and renewal of precursors for syntheses – e.g. chlorophyll). Other significant
functions include reproductive (type of leaves called sporophylls –
microsporophylls and megasporophylls and flower phylloms) and regulation
functions (phytohormones production). The most frequent, typical leaf is modified in
order to optimally work at material exchange with the environment, and this is why its
exterior as well as interior surface is large.
Besides, leaves can take over other completely different functions, which makes their
structure completely modified, e.g. defence function by the modification into spines.
Leaves are the most variable organ in an individual which ensues from its
numerous functions and necessity to structurally react to different environments in a
tree crown (light, wind, air humidity, temperature gradients). If a leaf completely
changes its appearance and structure not as a result of the photosynthetic function,
we call this change a metamorphosis. If it only reacts by slight deviations in the
structure or shape to exogenous impulses in order to optimally fulfil the function it is
called a modification.

6.1 Phyllogenesis – the evolution of a leaf

Leaf of vascular plants originated in a set of terminal plant body elements, sterile
telomes of psilophyton plants. Telomes used to planate, flatten, adhere and develop
in two directions. This is how large flat assimilation leaves, called macrophyll, of
present ferns and angiosperms were formed. A microphyll (a needle or scaly leaves
of angiosperms and meiophylls, thin single-vessel leaves of horsetails) is thought to
have originated by the reduction and condensation of macrophylls or their elements.
A flat leaf, found in Ginkgo biloba (i.e. among angiosperms) evolved from grafted
needles. Broad leaves of Japanese umbrella-pine (Sciadopitys verticillata) evolved
the same way.

6.2 Ontogenesis – the individual development of a leaf

A leaf develops on the apex tip as a leaf primordium. Further on, an axil bud
develops in its axil (scattered in angiosperms, regular in dicots). Buds of some woody
species (early flowering, e.g. Acer, Forsythia, Magnolia) divided by their origination
into leaf buds (bearing leaf elements) and flower buds (bearing flower elements);
buds in other species are mixed – first leaves grow, followed by flowers
(inflorescence – e.g. Tilia) on the tip of the shoot.
In the mild climate of Central Europe, buds of nearly all woods are protected by
covering scales. Naked buds (without covering scales) can be found in Frangula
and Viburnum lantana, semi-naked buds, covered only at the bottom side, can be
found in elderry bush (Sambucus nigra). Submerged buds (covered only at the
bottom of a leafstalk) are typical for Platanus. Scales that cover buds are formed in
various ways (opposite, double-line, spiral, sloping), they are either bald or
lashed, hairy, feltlike or veined (Aesculus).

51
The placing of a leaf in a bud prior to its opening is called the vernation. Woods may
have a flat vernation – front sides of leaves are one on top of another (Viscum); or
compound – if young leaves are folded in halves, each covered with the previous
one (Prunus); cirrus, where the blade is folded in a cirrus matter along the main
vessels (Fagus, Carpinus); or coiled (under or above). Besides the vernation,
estivation is classified too. Estivation is the relation between leaves in a single bud
which can touch each other, be lied one on anther etc.
Leaves on the stem develop acropetally, i.e. from the stem base with the oldest
leaves towards the apex with the youngest leaves. The interval between the
origination of one leaf and the origination of the next is known as plastochron.
Regarding the evolutionarily older plants such as turf, cycas, a leaf grows by
dividing of apex meristems. This means that the apex part is the youngest, spiral,
leaf base is oldest, develops first. The opposite process applies to conifers and
gymnosperms whose leaves grow by dividing of basal meristems – the oldest
parts are leaf tips. In spring, leaf tips get damaged by late frost or salty solutions
from urban and road salt treatment. In autumn, leaf tips and edges are the first to
mature and die. A budding and growing leaf is dependent on ready-made
photosynthates, it becomes autotrophic only after maturing, i.e. about two thirds of its
final weight.
Considering deciduous trees, low positioned buds on the base of a crown grow
first in spring, and buds on the crown top open last. There is the opposite process
in gymnosperms. It corresponds to the intensity of the upward pressure in roots,
which is very low in conifers. Opening of buds is affected by temperature and
lengthening of the daytime. In lower altitudes or on warmer slopes, woods of the
same species open buds earlier than in the mountains or on northern slopes.
Temperature is most important and is valued as a sum of effective temperatures.
Lower temperature requirements can be seen with mountain ecotypes of homoxylic
or scattered porose wood types. Round porose, for instance ash tree, prevent from
late frost by later bud development.
Lifetime and time of full functional capacity of leaves depends on a species and the
environment. In case of deciduous trees with falling leaves, leaves are functional for
one growing season, in case of evergreen deciduous, they persist for two or more
years (Hedera, Lonicera). Conifers may bear needles from 2–3 years (Pinus
silvestris), up to as many as 20 years (Pinus aristata). In general, lifetime of foliage is
shortened due to inadequate conditions that are either permanent such as
compacted soil, frequent or temporary, such as a fluctuation in humidity, temperature
etc. Leaves on the tree crowns margin change colour first, while basal and interior
more protected leaves grow old later (even though they develop buds earlier).
Maturing of leaves slow down by night light. This means that the length of a growing
season (defined by the number of days with favourable temperatures from spring to
autumn) within one crown may be even a month longer in the crown interior.
The autumn change in colour of leaves is related to the disintegration of
chlorophyll and covering of chlorophyll by carotenoids (yellow or orange leaves)
and antocyans (purple leaves, such as Parthenocissus inserta). Changing of colour
is a genetically controlled process during which the intensity of respiration rises,
energy rich substances, but also mobile nutrients such as phosphor or potassium
withdraw from leaves to storage tissues of stems, branches or roots. This process
mainly takes place at cold nights when day gets shorter; as it requires sufficient
humidity. The period of colour changes lasts about two or three weeks. At the
beginning of winter, leaves of deciduous trees fall off. This is how a tree gets rid of

52
waste matter of old, non-functional leaves. The process is controlled by growth
inhibitors and can be active due to a layer of parenchyma tissue of a leafstalk base,
whose solidity is violated by slimming of intracellular material, or its freezing, tearing
or dying off. After tracheids are torn, a leaf falls off. When a leaf is off, a new wound
may be either covered by a protective cork layer. The fallen foliage protects the
soil from drying, enables the nutrition to soil organisms, and it generally gives
organic and mineral materials back into the soil.
Though, needles of some conifers and leaves of some greenhorns (Hedera helix)
change colour in autumn, as well, without a follow up fall off. Their colour is
affected by the change in light intensity, day length and lowering temperature.
Needles of spruce and pine damaged by winter drought are redish, their fall off is
delayed. Insufficient nitrogen, magnesium, iron etc. nutrition leads to yellowing of
leaves and may be caused by the lack of nutrients in the soil or limited root activity.
We can sometimes see that a part of a tree foliage is green and another part is
yellowish. This may mean that conductive pathways of a branch may be
mechanically (e.g. by breaking) or biotically (e.g. fungi infection) damaged. Leaves on
such a branch have a worsened nutrition and water balance. Brownish colour of
leaves may be caused by the replacement of the central atom of chlorophyll (e.g.
magnesium) by an atom of hydrogen (in acid conditions, i.e. if vacuoles are
damaged, due to foliage decomposition or due to acid rain). If there is a critical lack
of water, trees plan to decrease transpiration area by premature fall off of leaves.

6.3 The arrangement of leaves on a stem

A spot where a leaf touches the stem (as part of the node) is called a leaf insertion;
and after a leaf falls off, this spot is called a leaf scar. In the leaf scar, you can
observe leaf tracks (spots after vascular bundles) and sometimes even pillows. The
shape and size of a scar and track character varied largely and become
characteristics for discerning woody species in the winter period. Regarding the
position on the stem, there are single leaves in a spiral (alternate), or pairs of leaves
on one node – opposite or whorled.
Shaded branches could have an alternate distribution of leaves transforms into
horizontally placed branches that are secondarily distichal, organized in one plane
(Tilia, Fagus, Picea). We could distinguish leaves by the angle between leaves and a
stem as upright (angle less than 45° degerees) and protruding (45° to 90° degrees).
The opposite position of leaf pairs facing each other may be cross, i.e. in two
vertical mutually orthogonal planes (often on upright branches). Even opposite
leaves may change into distichal. More than two leaves may grow from one node,
such as in Catalpa.
A leaf occurs in a plant in five categories: cotyledon in embryo, primary leaves in
a juvenile stage of a plant development (sometimes with shapes deviated from typical
trophophylls) and assimilation leaves (Fig. 24 and 25). In the flower part, there are
upper bracts (bracteoles) and flower phylloms. On the basis of a shoot, there are
cataphyllas presenting a shift from bud bracts to assimilation leaves of a growing
shoot. They may be scaly, tiny, their axel buds are small, undeveloped; they grow as
spare primordia and they develop branch collar in old branches.

53
Fig. 24 Germinated beech seedlings plotted above the soil surface cotyledon leaves
(epigeic germination) which become green (chloroplast development) and start
assimilation.

Fig. 25 Germinated oak seedlings plotted above the soil surface leaves (hypogeic
germination). Cotyledon leaves lie on the soil surface a support root and stem
growth.

6.4 The outer structure of assimilation leaves

Assimilation leaves of gymnospermous woods are made of a blade (lamina), a

petiole and a sheath. Any of these parts may be absent or become stunted and their
roles may be taken over by stem (Sarothamnus). A blade is a flattened part of a leaf.
It is usually green, but sometimes can be stained with green, yellow or white spots.
Some leaves can be red due to antocyans. Some decorative conifers have a delayed
generation of chlorophylls – and the light colour of young needles contrasts older
green ones (e.g. Picea abies `Argenteospicata`).
A blade bears typical characteristics of the species (shape of the outline, edge
and base, type of outline, trichomes, venation etc.) A simple blade is either undivided
or subdivided by lobes of various depths. They are arranged in various patterns:
54
pinnately or palmately (lobed into 1/3, 1/2, or 2/3, almost reaching the main vein or
blade base). If a blade is completely divided into individual leaves, the leaf is
compound – pinnate (odd-pinnate, even-pinnate, brake-pinnate, multi-pinnate etc.)
or palmate (tripalmate or multipalmate). In pinnately compound leaves, the main
spindle (or side spindles) is held by yokes of leaves (leaves facing opposite).
The rate between the perimeter and area of a leaf blade is often higher in
xerophytes than in sciophytes or hygrophytes (it is related to the resistance of the
bordering layer of air above a leaf that prevents from an excessive loss of water). The
tolerance of leaves to solar radiation and dryness is related to the consistence of
a leaf blade and its reduction. Sclerophylls are solid, hard, tough and resilient
leaves. Soft blades (malacophyllas) reduce their resilience by trichomes, succulent
leaves increase their volume (water storage tissue).
Leaf blades are mainly symmetric; however, in some woods (Ulmus, Fagus, Morus)
there is a certain asymmetry grounded as early as in the bud. For instance, in
horizontal branches of elm, the inner half of a leaf towards the branch apex is larger
than the outer half, a blade extends at the bottom near the main vein. An opposite
asymmetry is typical for beech – the largest is the outer side of a leaf. The leaf
heterogeneity means various behaviour and features not only of its right and left
halves, but also their marginal, central, apical and basal sections.
Shaded, horizontally arranged branches have a leaf mosaic (Fig. 26). This means
heterophylly, different length and angle of petioles along with various sizes of leaf
blades. This ideally utilizes the area and the leaves may optimally use the diffusion
radiation. There is also a special case of heterophylly when a plant has differently
formed assimilation leaves (Fig. 27). An individual plant bears leaves of the same
category of different shapes (e.g. lobate pinnate sterile leaves and compound leaves
bearing flower shoots Hedera helix).

Fig. 26 Leaf mosaic develops under low light condition by leaf blade and petiole
length changes. This arrangement enables maximal light capture.

55
Fig. 27 Heterophylly means that different leave shapes could be found on the one
individual.

A sheath is formed by an excessive development of a base part of a leaf and is

typical for monocotyledons – e.g. dracenae, banana tree, calla etc. Leaf sheath
grows around the stem above a node as high as another node, so the stem is
sometimes hidden. Leaf sheaths may take part in the mechanical stability of false
trunks.
A petiole is a stem part of a leaf that connects a leaf with a stem. Actually, it
originates in a bud, but its growth is delayed compared to a blade development.
Lower leaves of the upright shoots often have longer petioles than the upper leaves
(Acer). A petiole may turn leaf towards the optimal radiation (in case of needles,
e.g. spruce, they are repositioned in a bud in a spindle, into the open position of a
crown top or distichal in shade); the length and angle of petioles are also utilized in
the formation of a leaf mosaic. Flattened petioles increase leaf mobility, which also
helps increase water evaporation (Populus tremula). Sometimes, the petiole
extends in a hood preventing a bud during its development (Platanus). A leaf with
a well developed petiole is called a petiolated leaf, a leaf without a petiole is called
sessile leaves. We can further distinguish the following means of a leaf attachment
to a stem:, sessile, clasping, connate (the bases of two opposite leaves are joined),
sheathing (lobes are joined around the stem). If a blade is lacking, the stalk may
expand to phyllodium (Acacia, Mimosa). Stipules often replace a sheath on both
sides of a stalk. They are flat, scaly or leafy, either attached to the stalk only on its
base or grown up together on both sides. Stipules (membrane, scaly) protect (leaf or
flower) buds and soon fall off or change into thorns occasionally (see.
metamorphoses).

6.5 The inner leaf structure

The leaf is covered by the epidermis with a cuticle, stomata placed on the bottom
of a leaf (in hypostomatic leaves – in majority of local European deciduous leaves)
or on both leaf sides (amphistomatic leaves– Salix alba, Syringa). The leaf surface
may be smooth (matte, shiny, depending on the wax cuticle structure), wavy,
concave, with frequent trichomes. The epidermis is usually formed by a single layer
of cells. If multilayer (in sunny needles), then deeper cells are called the

56
hypodermis. The inner space of a leaf is filled with the primary tissue mesophyll.
There is a network of vascular bundles with colenchymatic or sclerenchymatic
sheath, i.e. venation (conductive and supporting system). Sheaths of vascular
bundles in woody plants leaves (except for e.g. hydrangea with isobaric leaves)
reach both the lower and upper epidermis. They are divided into more or less
independent alveolas (spaces), known as heterobarical (Fig. 28). In case of an
injury (e.g. insect sting or bite), the affected group of alveolas dies and the
surrounding ones remain functional because the pressure changes do not reach
them.

Fig. 28 Example of heterobarical leaf. Upper panel – Leaf damage is visible only in
the few alveolas and neighbouring alveolas are only partly damage. Lower panel –
cross-section through middle vein – sclerenchyma sheath which connect upper and
lower epidermis is visible.
Leaves are protected from insulation damage by turning their edges towards the
sunlight (Fagus). A similar reaction is observable with chloroplasts in individual cells
– cytoskeleton helps them get positioned so that they could take in as much solar
radiation as possible in the weak sunlight, and as little radiation as possible in the
direct light. According to the degree of tolerance to density of radiation, leaves in tree
tops are classified into sun or shade type (Fig. 29), differing by the width of cuticle,
epidermis, number of mesophyll layers, density of stomata, trichomes etc. (sun leaf
has a thicker skin, cuticle, thick and multilayer palisade parenchyma, a higher density
of stomata, a little volume of intercellulars and a higher density of venation; generally,
higher weight of dry matter related to a unit of leaf area). The position of a leaf in
space defines its water saturation – if deficient, they fade, or even die in a green
state.

57
Fig. 29 Example of sun shoot with dense needle organization and thicker needles
(upper panel) and shade shoot with spare needle oraganization

Mesophyll is a basic leaf tissue derived from the stem primary bark. It fulfils the
assimilation – photosynthetic function (chlorenchym). In bifacial leaves it is divided
into palisade and spongy layers of parenchyma (Fig. 28 and 30). On other hand,
this distinction is less obvious in the monofacial leaves, similarly as in the case of
needles of angiosperms (whose mesophyll makes shoulder cells – spruce, pine. In
yew and fir needles, mesophyll is divided into the palisade and spongy parenchyma,
and darker upper and lighter lower sides are well visible. Plants with C4
photosynthesis have a parenchyma sheath around vascular bundles (coronary
type). What is important for the exchange of gases is segmentation of mesophyll –
size of intercellulars 25–50 % of the total leaf volume and the surface area of
mesophyll cells (i.e. inner surface that may exceed 10–20 times the outer surface of
a leaf – it serves for speeded gas diffusion and solution of gases in water film of cell
walls).

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Fig. 30 Inner structure of bifacial leaf – consist of palisade and spongy parenchyma

A cuticle is a layer produced by epidermis walls on the outside of above ground

structures – leaves and stems. It is made of wax. These are esters of higher
monocarboxyl acids and higher aliphatic alcohols, usually in compound with
paraffines, alcohols, etc. The cuticle prevents from drying, but also from an
exchange of oxygen and carbon dioxide. It is typically structured (into trichytes,
lathes, grains, moulds etc.), however, urban atmosphere, polluted by exhaust gases,
aggressive aerosoles and emissions lead to its damage. At the end of a leaf life, it
is discontinuous and crackly, which may make cuticle transpiration exceed
stomata transpiration. The rate of dying of leaves is enhanced by water stress.
A stoma consists of two guard kidney-shaped cells with an intercellular space in
between; outer walls of both cells are fixed to the surrounding epidermis cells. The
openness of the stoma can be regulated. While surrounding epidermis cells do not
contain chloroplasts, the guard stoma cells contain them. Due to the
photosynthesis, assimilation of saccharides and potassium ions, stomata can
control water accumulation, which helps them change the volume. The utilized size of
a cell wall makes them long if they are saturated with water, which opens the stoma.
On the contrary, if there is a lack of water, cells weaken, get shorter and the stoma
shuts. Stomata are typical for the leaf epidermis, but they also exists in the epidermis
of stalks and young stems. Open stomata supply chlorenchym photosynthesis with
carbon dioxide and cool leaves by water vapour (transpiration). Moves of guard
cells are driven by the temperature, light, water content in leaves and the soil, CO2
content in intercellulars of a leaf, wind etc. The rate of gas diffusion depends on the
openness, size, shape, position and density of stomata; generally, on stomata
resistance. A high stomata resistance can be observed in leaves with
submerged, tiny, densely grouped (800–1000 units per mm-2 ) and quickly
reacting stomata protected by waxes and trichomes.
Hydatods, made of two guide cells with are permanently open stomata. They are
located on leaf tips or on the edges of blade teeth or fringes. A tissue with numerous
intercellular spaces leads to them (epithem). They serve for gutation, i.e. exclusion
of excess water from leaves (Salix, Ulmus, Sambucus). There are also wood species
that lack gutation and hydatods (Tilia, Acer pseudoplatanus etc.).
Trichomes are unicellular or multicellular, simple or branched outgrowths of
epidermis cells of leaves and stems. The simplest, papillae, cause the velvet
appearance of petals. They enable easy capturing of pollen on stigmas. Unicellular
trichomes may function as water reservoirs, plant protection from excessive water
vapour, radiation, changes in temperature etc. Branched or multicellular trichomes
make a perfect isolation layer (Hippophae, Viburnum lantana). Some types are
important for their secretion activity (secretion trichomes).
A file of vascular bundles in a blade of assimilation leaves and other leaf organs
(e.g. petals) is generally called venation. It is sometimes more visible from the
bottom of leaves, and if leaves are transparent, they can be seen against the light.
The main veins in blades with palmate venation direct in rays from the tip of a stalk
(or blade base, such as Acer platanoides); in pinnate, side veins (first class) turn
from the main vein (in the long axis of a leaf) and these branch further on into veins
of higher classes (Fagus, Carpinus etc.). In monocots, venation is parallel, rows of
veins of similar width direct from the base to the tip of leaves (bamboo), it is often
also pinnate, e.g. banana and palm trees. Dichotomous venation of tree ferns, or
ginkgo leaves divides several times into halves on the way from the base to the tip of

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a blade; it is also an open venation. Gymnosperms have veins connected with fine
anastomoses, whose venation is closed.
6.6 The inner needle structure

The structure of needles must allow them to live a longer life and to overcome a
relatively long season of low temperatures or frost in winter and, at the same time, to
survive heats and droughts and to be functional in the growing season. This is why
needles have a xerophyll structure. Needles are grounded in buds of the previous
growing season which affects their future number and structure. They begin
growing in early spring of the next year, (depending on nutrients and photosynthates
from older years) and they attain their permanent structure in two weeks time. As
they grow old, needles change into a shade type structure, which is reflected in the
pigment fond. Their structure is really close to the structure of a stem. On the surface,
there is epidermis (with extra cuticle made of wax layers or fine trichomes), below
which is an inner layer, hypodermis. Shoulder-like cells of mesophyll are often
homogenous and correspond to cells of stem primary bark; there are frequent resin
ducts.
The boundary between the mesophyll and the central cylinder is often made of an
endodermis. Lying tracheids and protein cells of transfusion tissue are placed
below the endoderm. This tissue is in contact with the central part – collaterally
arranged vascular bundles only divided by very thin phloem-xylem rays. The xylem
part is close to the upper (adaxial) side of a needle, the phloem part to the lower
part (abaxial).
Procambium in needles separates more sieve cells than tracheids. The shape of the
needles cross section may be affected by the number of brachyblast needles (Pinus),
rate of radiation during needle growth (Picea) and the arrangement of mesophyll
(Taxus, Abies, Larix). Needles are sensitive to sudden changes such as
disposition to light, wind, temperature fluctuations. If trees cannot take in enough
water for frost-proof modifications before winter, they suffer from water stress and
photooxidation of chlorophylls. Winter and spring are not favourable for them,
especially for the individuals living near roads treated with salt. Needles, and young
needles in particular, could get damaged due to frost drying.

Fig. 31 Cross-section of pine needle. Stomata are nested and arranged in lines.

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6.7 Evaluation of foliage state

The relation between foliage and other woody plant structures, especially
heterotrophic, is a widely used diagnosis means of a woody plant prosperity. E.g.
relative weight of foliage expresses the relation of the weight of dry leaves mass to
the total dry mass. In early stages of tree life, the dry matter of leaves makes up to
50% of the total mass, and due to storing organic matter into roots, trunks and
branches, this rate falls down to 5% or even below 1% in mature and old age. The
density of foliage, utilized space of leaves in other words, expresses the volume of
a tree crown space per an average leaf – which is higher in dense crowns of young
trees than in less dense crowns.
The leaf area index expressed by the rate of the total cumulative area of foliage
related to the area of crown projection on the ground. It can successfully express the
area of reception of the solar radiation as well as the area of plant surface. This value
varies from 3 to 5 in large deciduous trees (i.e. 1 sq metre of soil surface is
shadowed by 3–5 sq metres of leaves), and may reach as much as threefold in case
of conifers.

6.8 Leaf metamorphoses

In general, in the course of the long evolution of plants, there have been many cases
of complete metamorphoses of the primary functions of leaves accompanied with
changes in the structure. Cotyledons, lower and upper bracts (braceae – supporting
leaves in whose axil a flower or inflorescence develop on a stalk). Other new
structures include covering scales of buds or conifer cones, supporting scales of
catkins that further lignify (Alnus) and covers of inflorescence (Cornus mas).
Metamorphoses of woody plant leaves also include shoots; spines that have
originated from the whole leaf (Berberis) or from stipules (Robinia). And, finally, a
flower originates in the whole complex of leaves metamorphed for the function of
sexual reproduction.

7 Propagation and reproduction of woody plants

Woody species are perennial plants anchored in the soil by the root system.
Therefore, any move from one place to another for finding a shelter is not supposable
and an individual has to be able to conform to all changes on its site by its own
biochemical, structural and growth activities. There are only exceptional cases when
a plant could move to another position, e.g. due to the solifluction, landslide (mainly
on slide rock), water flooding or moving of the soil by water or wind erosion, moving
by snowslip, falling of banks by water abrasion etc. Nonetheless, as these plant
movements are driven by exogenous factors, a plant has not got much space for the
acclimatization and it is a rare phenomenon if a plant survives.
Yet, there exists a movement of woody plants, i.e. migration. Species spread to
new sites that are more favourable to them (immigrate); they could even leave their
present sites which are no more suitable (emigrate). This movement is facilitated due
to regularly produced diasporas. Spreading of diasporas from the parent plant
(dissemination and propagation) into various distances is defined by their shape,
weight, character, number and type of activity of the spreading force. This force

61
includes a presence and movement of animals, direction and speed of wind, water
etc. For instance, birch (Betula) anchenes get carried about 90 metres away, ash
(Fraxinus excelsior) about 2 metres, spruce (Picea abies) seeds about 6 metres.
7.1 Means of reproduction and types of diasporas

A diaspora may have a generative origin (changes in the genetics, i.e. by means of
seeds or fruits – acorn e.g. Quercus, Fagus, berries– Berberis etc.), this way of
spreading is called a dissemination. A group of individuals that originate in the
genetic way is called a generation.
A diaspora may even be of an asexual origin – vegetative. This a direct
continuation of a plant life via its body parts – via shoots (e.g. Robinia pseudoacacia,
Populus tremula, Rosa), via branches that easily break off and then root (Salix,
Populus) or stem or trunk stools of fallen off individuals (Quercus, Tilia, Salix) or via
basal branches creeping on the ground that gradually root (Picea, Fagus, Pinus
mugo and the like).
Spreading of vegetative diasporas is generally called a propagation. It is mainly
practicable in species with high regeneration ability. The vegetative propagation of
trees is can be very successful in unfavourable conditions; it diminishes the death
rate of new individuals. This is thanks to the fact that the little plant remains
connected to its mother plant for a longer time, and the polycormon may be more
successful in the competition with the neighbouring plants. Propagation units could
even settle sites with dense vegetation where seeds would have problem germinating
and new sprouts would have problem developing. The vegetative reproduction is a
reserve way of keeping a species on a site if conditions for the sexual reproduction
(flowering) are not favourable (e.g. low temperatures in the mountainous altitudes
etc.). New individuals originating in an asexual way make a clone.

7.1.1 Artificial vegetative reproduction

Human spreads and retains interesting cultivars, variants, hybrids etc. in both the
ways – in an autovegetative way (cuttings, slips, layering, offsets, meristem cultures
etc.) as well as in a heterovegetative way (grafting – suitability of a graft and a stock
influencing each other must be taken into consideration).

These could be well applied to a number of cases, but always being aware of the fact
that this is a rare phenomenon in the phylogenesis (long-term plant evolution on
Earth). On the other hand, vegetative reproduction is typical for the reproduction
of lower organisms (e.g. bacteria) with very short lives; mutation and horizontal
transcription of information play a vital part in the survival of a species.
Artificial vegetative reproduction of woody plants is only reasonable in special cases
and for special intentions. For instance, if we wish to sustain multiple or distant cross-
breeds, mutants or other genotype valuable individuals with a weak reproduction
activity, or senescent or senile individuals or if we wish to retain genetic sources and
create cultivation populations (testing plantations or populations for hybridisation
activities) and following populations for the mass reproduction.
Individuals in a clone may have a required united appearance and characteristics of
the parent organism, but they could also gain undesirable changes, e.g. direction of
growing and branching below the apex control value (especially conifers). In the adult
age, the expected height might not be reached, there may be irregularities in the
formation of generative organs; most frequently with monoecious species whose

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flower sexuality is not controlled by heterochromosomes (e.g. grafts of fir tend not to
reproduce female cones). Furthermore, they could get old sooner, their stability could
be worsened due to lower wood quality and a persistence to phytofagous insects and
phytopatogenous fungi. Another disadvantage in the changing environment is a
lowered adaptability due to which cloned individuals need to be regularly observed
and treated (e.g. limiting of suckers on drafts).

7.2 Plant life cycle and the significance of seeds in phylogenesis of

plants

The plant life cycle consists of sexual (generative, haploid) and asexual (vegetative,
diploid) generations. In the phylogenesis, there is an evident shift from completely
absent changes of reproductive organs in microorganisms and rare occurrence with
fungi to well developed, multiple and regular changes with an independent (algae –
green, brown, red) and prevailing gametophyte (mainly in bryophytes and
pteridophytes).
Seed plants have asexual diploid sporophyte called zygote. A seed develops from
the zygote, and finally an autonomous plant consisting of a root, stem, leaf; sexual
haploid gametophyte is reduced to a dependent part of sporophyte and is created
only in an adult stage, i.e. after the first flowering (male in a pollen grain, female in
an ovum). Plant life cycle enables for a regular change of genetic information
when forming a new generation (by combining and recombining endowments) and
achieving a longer life of an individual.
Seed plants present three wide taxonomic groups. One of the groups, angiosperm,
have taken the greatest steps in the plant evolution – a new type of seeds developing
after a double fertilization of an ovum (one pollen grain nucleus merges with the
ovum cell nucleus to form a zygote; an embryo of a new individual develops from the
zygote; the second nucleus of the pollen grain merges with the central nucleus of the
embryonic cover and becomes a ground for the nutrient tissues – endosperm whose
cells have a triple chromosome code). This background, which was given to a
daughter generation by the parent generation, resulted in a swift entrance of
gymnosperms, development of a huge number of flora species, and a quick
occupation of all the continents in late Mesozoic era and in early Tertiary era.
The formation of seeds and their number is only possible in maturity, i.e. fertility of an
individual, and is affected by a number of photosynthates, rate of these
photosynthates transported to the generative organ and the size of seeds into which
energy is transformed. The high energetic demands of flowering, development of
fruits and seeds do not allow many tree species a regular, yearly activity, but only in
seed years which may be of various period depending on a species, and
endogenous and exogenous conditions. At present, when the total year temperatures
sum grows, some species flower more frequently which is not suitable for them. A
number of individuals (mainly in the Picea family) flower every two or three years.
Flowering (and mainly formation of female cones) takes place in the top third of tree
crowns. Needles are more exposed to the solar radiation (and temperature changes),
higher transpiration (and frequent water stress) and higher demands on good working
photosynthesis. The combination of stressors may result in the dying of needles (and
creation of a hole) or in the limitation of elongation of growth and branching of the first
class (sudden thinning of a crown). It can also be the means of growing plants in
nurseries that cause changes in tree performance. A very sensitive species is larch

63
(Larix), which can flower as early as at the age of 8 years after having repetitively
injured roots.
Comparisons between species prove that the number of produced seeds is often in a
negative correlation with their size (a large number of very light single seed fruits of
birches, on the contrary palm tree Lodoicea seychellarum makes a fruit with the
largest seed in the plant realm weighing 1.8 to 27 kg). But in general, seeds are very
little variable, i.e. their special and weigh characteristics vary only slightly. Moving of
diasporas to new sites also depends on the distance from the source population, on
barriers that may prevent diasporas from spreading (mountains, waters etc.).
The number of life diasporas, germinated seeds, varies by species and
environmental conditions. The longest lifetime and germinability are observed in
seeds of annuals and biennials plants (e.g. goosefoot – Chenopodium album –
spurry – Spergula arvensis – up to 1600 years), while a relative short life is seen in
large seeds of e.g. walnut (Juglans regia), haselnut (Corylus colurna) or oak
(Quercus petraea), a very short lifetime is observed in willow (Salix caprea only
about 14 days).
The plant ability to survive in a dormant state in the form of diasporas is a result of a
long plant evolution and adaptations to exogenous conditions. In immobile plants
dormation of diasporas in connection with their dissemination is analogic to an
escape of animals from unsuitable locations. The length of dormation and conditions
of its interruption may be genetically defined (primary dormation) in some species
and present a permanent adaptation to climatic conditions in the centre of their
habitat.
A dormation period of seeds may be even caused by unfavourable exogenous
conditions (secondary dormation, quiescence). When conditions improve (e.g. in
species living in cleanings after coverings of the soil surface), seeds soon germinate.
Not only seeds and other types of diasporas, but also grown seedlings may remain
in some state of quiescence with a very limited metabolism during bad living
conditions. Under an adult stand of trees and bushes or in the shadow of a building,
groups of seedlings of woody plants could persist for a number of years (Fagus,
Quercus, Acer). Slow growing due to the weak radiation, lack of soil humidity (mainly
as an outcome of adult trees roots suction) keeps these groups of seedlings in a
juvenile stage of the ontogenesis with a relatively low yearly mortality. These are
"candidates" for a quick launch of growth as soon as the area gets cleared up.

7.2.1 Dissemination by diasporas of a generative type

During the sexual reproduction (i.e. creation of diasporas of generative origin,

seeds, fruits etc.) genotypes are renewed or completely new genotypes originate
during the formation of zygotes. This means that the variability among individuals
in a population increases. A hard competitive natural selection comes to the
scene (mainly with utilized adaptation characteristics) both in parents and young
population. The natural selection, affected by accidental events, isolation etc., takes
place under the pressure for the initiation and differentiation of flower organs,
formation of gametes, transfer of pollen and the follow-up germination, fertilization
and development of seeds and embryos, transfer of diasporas, and further on under
the pressure of new conditions on germination of seeds, overcoming early and
juvenile life stages of individuals in the same generation. Sexual diasporas are
located in a flower, sometimes there are also groups of microsporophytas and
megasporophytas.

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A flower is a metamorphosed shoot, i.e. shortened stem (peduncle) and a file of
metamorphosed leaves that participate on the sexual reproduction and spreading of
seed plants (Fig. 32). A flower comprises of perianth which is divided into a
calyx/sepals (usually green) and a corolla/petals (coloured) or sometimes is
perianth undifferentiated (stamens). Sometimes a perianth may be absent (Salix,
Fraxinus). Sepals, petals and stamens are either free or connate. Hard calyx sepals
protect the inner parts of a flower, mainly in the budding stage. It usually falls off after
blooming. On the inner bottom, there may be nectaries, e.g. in a lime. There are
cases when the calyx supplements the corolla and takes the corolla colour (Calluna).
The inner whorl, corolla, protects reproductive organs and attracts insects by its
colour, which is mixed from dyes diluted in a cell liquid or dyes in plastides. Lower
parts of its petals contain honey containers. Like the calyx, it is typically regular
(Tilia) or symmetric (Vicia family). In the Vicia family, it comprises of an upper pavis,
two lateral wings and two bottom leaves. Peduncle is typically quite small, it either
expands to a sheath or sometimes may be conical (Rubus), convex (Cerasus) or
extend and conves in a cupulla embrasing a flower (Quercus). The scyphus or an
exciple lower part of a flower, formed by the fusion of a peduncle with the lower parts
of a flower whorl and filaments of stamens, which embraces a fruit completely or
partly, is called a hypanthium.

Fig. 32 Flower structure. Bottom parts of petals and sepals and stamens merge and
form hypanthium. Receptacle is the modified or expanded portion of the stem that
bears the organs of a single flower.
Evolutionarily older species have spiral or spirocyclic flowers (position of flower
part in the spiral and in the circle), regular (symmetric by more planes) or bisymmetric
(symmetric by two planes). Flower parts of evolutionarily younger species are
arranged in circles (cyclical) that prone to be asymmetric (zygomorphic, symmetric
by one plane or asymmetric).
The floral diagram demonstrates the scheme of a number and position of flower
organs. It may be pictorial, which draws individual parts of a flower in a projection or
written, which expresses the floral diagram by means of formulas. The latter is often
applied in botanical encyclopaedias for determining plant species.
An inflorescence is a group of flowers arranged in a specific pattern on the common
main stem (a peduncle). Leaves are metamophosed into bracts (or suppressed,
reduced or absent, aborted). The main stem branches into side branches (rachis
or stalks – pedicals) of the first, second or higher classes. If the main stem remains
the thickest and longest among the other branches, this type is called the racemose

65
(indeterminate) inflorescence, blooming from the bottom upward, or from the
periphery to the centre – panicle, cluster, corymb, spike, spikelet, catkin, spadix,
umbel, composite plant. If side branches – rachis – are longer that the main stem –
peduncle, this is a cymose (determinate) inflorescence – blooming from the apex
downwards the basis, from the centre to the edge – cyme, little bunch, rhypidium,
dichasium, umbelliform, double cyme, bostryx, drepanium etc. Apex of simple
inflorescences main and lateral branches has flowers, while compound
inflorescences do not have flowers on the apex position. They use them to form
further inflorescences. If the inflorescences comprise of the same type
inflorescences, they are known as homoeothetic. For instance, they could be only
made either of cluster or cyme subunits. If the compound subunits are different, these
inflorescences are called heterothetic. These consist of both the cluster and cyme
units. From the perspective of evolution, inflorescence is more advanced that single
flowers.
Male reproductive organs are stamens, female reproductive organs are carpels.
Flowers with both these sexes are monoclinous (bisexual). Diclinous (unisexual)
are the ones that only have either male stamens or female carpels. If neither is
present, these flowers are sterile. Dioecious plants have separate sexes, i.e. only
male organs are formed in one individual and only female organs are formed in
another (Taxus baccata, Salix, Populus). Monoecious plants form flowers or
strobiles of separate sex (either male or female), but they are situated on the one
individual (Pinus, Populus). Polygamous plants have all male, female and bisexual
flowers (Fraxinus).
Stamens, with multiple pollen in gymnosperms, are arranged in a spiral into
strobili (cone). In angiosperms, stamens are divided into a filament and an
anther with four pollen sac with a connector. They are placed on the receptacle in
two or just one ring, they are altogether called androeceum. They produce huge
mass of pollen grains, mainly being of a round or eliptical shape and various sizes. A
pollen grain is a male spore of seed plants, it generates by the reduction division in
a male sporangium (anther). The pollen grains have two-layered walls – a tough
outer wall called the exine (with a cutin, sporopolenin) and a thin inner wall called the
intine. At pollen grain sides, there are often air-sacs (Pinus), in the Ericea family,
grains are grouped in fours. A mature pollen grain (in angiosperms) contains a
vegetative cell (tubular that disappears when pollen tubes germinate, this presents a
reduced procell, prothalium in bryophytes and pteridophytes) and a generative cell
(reproductive, giving rise to two generative nuclei in the pollen tube). There are
also the stamens that do not create pollen, and these could be modified into scales,
plates or honey containers.
Carpel, female reproductive leaves of gymnosperms are replaced by seed scales.
In angiosperms, they consits of a stigma, style and ovary bearing ovules. The set off
carpels is called gynoecium.
A stigma serves to capture pollen grains. It has various flaps, sculpturings and hairs.
Pollen in its mucous presence germinates in pollen tubes which penetrate through
styles towards an ovule. For instance, it takes place 2–3 weeks from the reception
of pollen to the germination in beech. The ovary makes the lower part of a carpel.
From the perspective of the position of stamens and perianth, we distinguish between
a superior ovary (other organs below ovary), half-inferior (other organs around the
upper part of an ovary) and inferior ovary (other organs above an ovary).
An ovulum is tied to the inner layer of an ovary by a funiculus that passes to the
ovule base (chalaza), where one or two integuments develop. They surround the

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inner multicellular tissue nucellus; with an opening on the top, which is a place
(micropyle) for an escape of a pollen tube in the process of fertilization. During an
ovule development, a megaspore differentiates from an inner nucellus under
micropyla, and this cell gives rise (either by dividing or directly) to an embryo sac.
After its reduction dividing, three of its daughter cells die, and the fourth,
megametophyte, presents a haploid female sporangium – embryo sac.
A female sporangium undergoes a multiple division in angiosperms. This results
in two groups of four cells; one group positioned near the micropylar and the other
near the chalaza ends. One cell of each of the groups differentiate a diploid nucleus
of embryo sac and a three-cell set remain (in micropyle an ovule set made of
oosphere and two helping cells – synergids; in chalaza, a triplet of cells are antipod).
One of two generative nuclei of a pollen grain fertilize the ovulum cell to form a
zygote. The other one fertilizes the central nucleus of an embryo sac to form a
triploid ground for endosperm.
A gymnosperm ovule is simpler. The embryo sac has a couple of archegonia at
the micropylar end where egg channel cell develops. Nucellus is a place of formation
of a haploid tissue, primary endosperm. This is formed by dividing a high number
of cells prior to fertilization. Ovules are naked, easily accessible to pollen grains. The
grains captured on a pollination drop released by micropylas are soaked inside as is
dries down. It germinates into a short pollen tube and releases a generative nucleus
that fertilizes only the oosphere after the entry into some of archegies.
The pollination is a transfer and transposition of pollen grains on a stigma or an
ovule micropyle. Pollen is transferred from one flower to another by insects
(entomogamous, tenomophilous species e.g. Tilia), by the wind (anemophilous, e.g.
Betula) or by animals (zoophilous, e.g. cactuses) etc.
Pollination if followed by fertilization, i.e. the fusion of male and female sexual
cells – gametes (male gametes are formed in a pollen tube, in which pollen grains
germinate after the reception on the stigma; female gamete – an ovum is placed in
an ovule) which results in the formation of a zygote. A zygote is a fertilized ovum cell,
the first diploid cell of a new organism originated in a generative way. In seed plants,
female gametes are larger. If they are immobile, they are called ovum. Male, smaller,
mobile cells are called spermatozoids (bryophytes, pteridophytes). Immobile cells are
called spermatic (in angiosperm and gymnosperms). A fertilized zygote grows into an
embryo, and an ovum develops into a seed, and an ovary changes into a fruit after
the fertilization.
Autogamy is a means of fertilization in which pollen is transferred onto a stigma or
ovum micropyles of the same flower as the pollen. Plants practising autogamy are
called self-compatible. In order to prevent from autogamy, stamens and pistils in a
flower mature at different times (dichogamy) in both sexes of woody plants. For
instance, in apples, stigmas mature first and they are fertilized from older flowers,
since their anthers open after stigmas have bloomed (proterogeny). In lime, the
principle is opposite – pistils mature after stamens have finished their fertilization
(proterandry). Fertilization by pollen of another plant of the same species is called
allogamy. An embryo may develop even without fertilization, by apomixis
(parthenogenesis, apogamy).
A seed is a reproductive element of angiosperms and gymnosperms that originates
from a fertilized ovulum (Fig. 33). Its surface is coated by testa (evelops from the
integument) and typically includes nutrient tissues – outer (perisperm from
nucellus) and inner (endosperm), sometimes only the endosperm. The testa is thin
and membranous (Juglans), leathery (Aesculus) or tough (Magnolia). The vital part of

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a seed is an embryo that may consume all nucellus and nearly all endosperm, while
the reserve matter is placed in its cotyledons (Fagaceae) or in a hypocotyl. Dicots
have two cotyledons, conifers have 2 to 13 cotyledons. A hilum may be visible on a
seed. A hilum is a scar after the detachment of a different colour than the rest of the
seed and with a trace of a vascular bound (Aesculus); a cicatris, little scar from a
micropyle opening (root base of an embryo is placed below), sometimes even a
raphe. On the surface of some seeds there are various formations serving for
spreading. E.g. a yew and spindle tree seeds include a colourful arillus made
during the fertilization.
An embryo is the youngest ontogenetic stage of a plant. The very first ontogenetic
stage is called pro-embryo, only containing a suspensor and an embryogene.
Generally, cotyledons, a hypocotyl, radicle and plumule develop. A plumule is a
foundation of the future stem and the first assimilation leaves. Germination, the
beginning of an embryo development into a young plant, is classified by different
hypocotyl growths into the above-ground (epigeic) and underground (hypogeic)
germination. The hypocotyl pushes plant cotelydons above the soil surface in the
epigeic germination (Tilia, Carpinus), while the growth of the hypocotyl in hypogeic
germination is inhibited and cotyledons remain inside a seed and an epicotyl
elongates (Quercus).

Fig. 33 Seed originates from a fertilized ovulum. In angiosperm, one generative

nuclei of a pollen grain fertilizes the central nucleus of an embryo sac to form a
triploid ground for endosperm. Testa evelops from the integuments.
A fruit (fructus) is a formation that includes seeds and was formed from an ovary
– this is a true fruit. The outer ovary wall develops into a pericarp. It is a cover of a
seed or seeds. By its position we distinguish between the outermost exocarp (oily
skin of various colours – cherry, plum, apricot), middle layer mesocarp – fleshy or
succulent parenchyma tissue typical for berries and stone fruits) and the inside
endocarp – hard stone with a seed hidden inside). By the quality of pericarp, we
classify dry and fleshy fruits. Dry ones are subdivided into dehiscent (vesicle, pod,
silique, little silique, capsule), in-dehiscent (nut, drupe, achene) (Fig. 34). The
fleshy fruits are subdividided into: pepo, hesperidium made of fleshy parts of the
endocarp. If other flower parts participate in a fruit formation, such fruits are called
accessory fruits (e.g. bottom parts of petals and sepals and stamens merge and

68
become fleshy and form hypanthium (Fig. 35); a case of which is rose-hip with
achene fruit, calyx becomes fleshy in mulberry etc.). False fruit is also a term for
gymnosperm species – cone, compound cone, conical berry (Juniperus), seed berry
(Taxus), seed drupe (Ginkgo). Fruits that develop without being fertilized, without
seeds are called parthenocarpic.
Simple fruits originate in a single flower and their seeds are closed in one ovary.
A group of fruits from single gynoecium joint by a receptacle or another type of
fusion within one flower form an aggregate fruit (e.g. aggregate vesicles in
magnolia, aggregate drope in blackberry). A group of free fruits is called a multiple
fruit – Ribes or variously joint or fused fruits (Fagus, Castanea, Lonicera), originating
from the whole inflorescence, i.e. is carried by a common stem.

Fig. 34 Achenes (true, dry and in-dehiscent fruit) of beech and oak develop in the
cupule which is of stem origin.

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Fig. 35 False fruit of rose. Seeds (achene) were formed from an ovary after ovulum
was fertilized. They are cover by hypanthium which get mostly reddish and attract
animals to eat them – endozoochory. There are dry sepals on the top of hypanthium.

The dispersal of reproductive units of plants, such as spores, seeds, fruits, fruit
groups etc. from the mother plant is called the diaspore dispersal. Based on the
factors utilized in the dispersal, the following types are distinguished: anemochory –
wind dispersal (Betula; Fig. 36); zoochory – via animals, this may be exozoochory
or epizoochory on body surface, e.g. dry diasporas with adhesive elements,
endozoochory – via ingestion e.g. fleshy and colourful diasporas, synzoochory –
esp. by birds that carry diasporas on a special purpose, e.g. jay dispersal of acorns;
antropochory – dispersal by humans purposefully or unintentionally, e.g. cultural
plants, decorative plants, jagged weed); hydrochory – dispersal by water, rain;
barachory – dispersal of heavier, round or oval diasporas that roll down slopes –
spruce, pine cones, oak acorns); autochory – self-dispersal by special mechanisms
including growth. The above classification is only schematic, many plant species
utilize combined ways of dispersal.

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Fig. 36 Betula pendula as an example of tree adapted to anemochory (seeds
dispersal by wind)

8 Basics of woody plant physiology

Scheme of main physiological functions of above- and below-ground is given in Fig.

37. Water and nutrients are taken up by roots from soil and transport through sap (S)
to aboveground system. Water, nutrients and CO2 taken from atmosphere run
photosynthesis in leaves together with radiation (PAR – photosynthetic active
radiation). Leaf heating and wind speed up water vapour from leaves – transpiration,

71
but at the same time CO2 a O2 are received by plant. Saccharides are transported
from leaves in the phloem (P) to tree apex, cambium (C), phelogen (F) and mainly to
roots. All living cells respirate i.e. consume saccharides a release CO2. Inner wood
consists of heartwood without physiological function.

Fig. 37 Scheme of main physiological functions of above- and below-ground

(according Požgaj et al. 1997)

8.1 Photosynthesis

Photosynthesis (from Greek phos, photós = light) is an anabolic synthetic action

occurring in autotrophic, photosynthetic organisms (photosynthetic bacteria, algae
and green plants), whose activity is necessary for the existence of all other
processes in life organisms, i.e. respiration and syntheses of all types of
organic material of the whole biosphere on Earth. Photosynthesis facilitates the
reception and binding of electromagnetic energy of solar radiation into the

72
energy of chemical bonds, its utilization for breaking water molecules (photolysis),
which enriches the atmosphere with oxygen and which facilitates incorporation
of atoms of carbon, oxygen (from atmospheric carbon dioxide) and hydrogen
(from water H2O) into simple saccharide molecules. Every mol of receipted carbon
dioxide corresponds to 477 kJ of potential energy.
By subsequent processes plants create small molecules of aminoacids, nucleotids
and fatty acids incorporated in various metabolic ways into proteins, nucleic acids,
polysaccharides and lipids, i.e. into the foundations of the above-molecular functional
and structural systems. However, these complex synthetic processes come after
the respiration, a process characteristic for all life organisms, i.e. their living
cells. Organic substances, primarily originated in photosynthesis, function as
nutrition molecules for heterotrophic organisms unable to live only from inorganic
sources (herbivores, omnivores, parasites, saprophytes). In this view, all
photosynthetic organisms are producers of organic matter and all other life forms –
consumers, reducers (living of decomposition, mineralization of died bodies of any
organisms) are dependent on them.
Photosynthesis involves photochemical processes requiring light (primary
processes), enzymatic processes not requiring light (i.e. dark, secondary
reactions) and diffusion processes facilitating the exchange of carbon dioxide
and oxygen.
Solar radiation is absorbed by photosynthetically active pigments that occur in leaf
mesophyll cells, in primary bark of petioles and stems, i.e. branches and other green
parts of the above ground system:
1. chlorophylls: higher plants contain chlorophyll a and chlorophyll b.
Chlorophylls are porphyrins, derivates or tetrapyrole joined to complexly bound
cation Mg2+.These are very similar to human haemoglobin in erythrocytes, but
containing a ferrous atom. Their colour and function is defined by the molecular
structure including the arrangement in the protein complex of chloroplast
membranes (thykaloids). The content of chlorophyll a, blue-green, more active, is
2–3 times higher than the content of yellow-green chlorophyll b in shaded places,
twice and more times in leaves in the sunlight. In dry matter of leaves,
chlorophylls make about 0,6 % to 1,2 % depending on a species, position in the
crown, development stage, radiation conditions, nutrition etc. A daily renewal of
chlorophylls is 5–8 % of the total content.
Porphyrin sections of chlorophyll molecules are bound to non-polar, linear chain
of phytol settling in a chloroplast membrane which secure them a special
position.
2. carotenoids: orange carotens (α- and β-caroten) and yellow and brown
xantophyls (xantophyl, zeaxanthin, violaxantin etc.). Carotenoids are isopren
derivates, however, they have a linear arrangement. The alternation of double
and triple bonds with delocalized free electrons is typical for them (as well as in
chlorophylls).

This makes both systems capable of receipting only a part of radiation of a

certain range of wavelength, the remaining radiation is reflected and released
back. This reflected radiation is perceived by human eyesight as a complementary
colour to the absorbed colour (e.g. leaves appear green, while they absorb blue and
red range of visible spectre). Carotenoids have light absorbing, antioxidation and
ecological (a warning signal for pollinators and animals that disperse seeds of eaten
fruit) functions.

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The first stage of the photosynthesis is the reception of radiant energy. To be more
specific, this means the entry of radiation influenced by spatial arrangement of
branches and the foliage area in a tree crown (or above ground system of a bush,
liana and other forms), arrangement of chlorenchyma (tissues with cells containing
chloroplasts in general), arrangement of chloroplasts within cells and arrangement
of pigment antenna systems.
The core reaction centre of photosynthesis is chlorophyll a. Other colours serve as
complementary colours, that, with the cooperation of enzymes and proteins form
special light reception antennas – these serve for the reception of photons and
control of their flow to chlorophyll a.
In the first stage (primary photosynthetic processes), the solar energy is captured
and stored temporarily into the energy of chemical bonds of special small molecules
of ATP adenosintriphosphate (in general, this molecule serves as a storage of quick
energy in all life cells, even animal cells). At the same time, energy is placed into a
molecule of a reduction agent – coenzyme nicotinamide adenine dinucleotide
phosphate (NADPH). Both molecules are primary products of photosynthesis and
serve as energy and reactive chemical groups transmitters. The processes of this
stage take place on thykaloid membranes of chloroplasts in two interconnected
steps by means of photosystems (complexes of protein molecules and hundreds of
molecules of photosynthetic pigments). This is the photosystem I, evolutionarily older,
existing already in photosynthesising bacteria. Then there is the photosystem II,
evolutionarily younger. Photosystem I absorbs radiant energy of wavelengths about
700 nm (reaction centre P-700) receipts hydrogen for the reduction of NADP;
photosystem II absorbs radiation of shorter wavelengths (reaction centre P-680),
dissolves water molecules resulting into a molecular oxygen and hydrogen, which is
used for ATP generation. Oxygen, as a by-product of primary processes is released
into the air and is necessary for breathing of all aerobic organisms (including
humans).
In the second stage (secondary photosynthetic processes), molecules from the
primary processes are utilized. They give energy and a reduction potential – i.e.
drive the process of carbon fixation. This produces saccharides out of gaseous
atmospheric carbon dioxide (CO2, entering plants through stomata or lenticels) and
water (H2O, taken in by plant roots from the soil). This process is located in
chloroplast stroma. It does not require light, but depends on the primary stage
anyway- it finishes soon after getting dark as well.
The process of incorporation of carbon from CO2 into organic bonds is defined
by a plant species. In case of woody plants, there are two types. In the Calvin cycle,
the acceptor of CO2 is 5-carbon saccharide ribulose 1,5 biphosphate RuBP (enzym
Rubisco), which reduces after the reception of a carbon atom into two three-carbon
molecules. This way is used by C3 plant species and carboxylation is catalysed by
specific enzyme Rubisco. In the other cycle of Hatch-Slack pathway, CO2 acceptor
is a tree-carbon (phosphoenolpyruvate; PEP), providing a 4-carbon product after
the fixation of a carbon atom. This is a type of photosynthesis in C4 plants where
both Rubisco enzyme and PEP carboxylase enzyme participate. The final products of
both synthetic ways are saccharides (photosynthates) – outcome compounds of all
other substances in body and a renewed receptor CO2 (i.e. 5 or 3 carbon acceptor –
Acc).

In central European conditions, plants with C3 photosynthesis prevail. They are more
sensitive to high temperatures, droughts and lack of nitrogen. Hatch-Slack

74
process is present in a range of thermophyll, e.g. tropical species. Casings around
vascular bundles with cells containing chloroplasts are typical for plants with the
Hatch-Slack process. However, there are even such cases of plants with both types –
C3 and C4 photosynthesis, such as corn in our climate.

Primary photosynthetic processes may be expressed by these entry factors

and outcome products:

radiant energy (8 photons) + chlorophylls + 2 NADP+ + 2 ADP + phosphate + 2 H2O

→
2 NADPH2 + 2 ATP + O2

Secondary processes utilize outcomes of the primary photosynthetic process

and renew the entry factors:

nCO2 + nAcc + 2n NADPH2 + 2n ATP → glucose +2n NADP+ + 2n ADP + 2n

phosphate + nAcc + thermal energy

Nonetheless, saccharide and ATP molecules (with rich energy phosphate bond
generated by photophosphorisation) are only utilized as a source of energy for
chemical bonds and a source of material for the preparation of a range of other
small organic molecules necessary for a plant cell.
8.1.1 The impact of exogenous and endogenous factors on
photosynthesis

The photosynthesis outcomes are saccharides (photosynthates), basic compounds of

all other material in body and a renewed acceptor of CO2. The quality of
photosynthesis in view to other life processes (respiration) is manifested by CO2
consumption (or release of O2, clean photosynthesis). The photosynthesis
capacity of a species (in a given stage of development and activity) means the
highest speed of reception of CO2 in a typical content of CO2 in the atmosphere and
optimal conditions of other factors (light, water, temperature, nutrients etc.). The
potential photosynthesis is defined in the conditions of optimal CO2 pressure. The
compensation point (expressed by radiation, CO2 availability, temperature...) is a
state of a woody plant or its part - branch, leaves..., with a zero exchange of CO2, i.e.
balanced rate of photosynthesis (consumption of CO2) and respiration (release of
CO2). If the conditions of the compensation point are long-term, a plant or its part
weakens and dies prematurely. In shaded woody plants this often results in a limited
growth, a limited immunity from pathogens and a loss of basal part of a crown (self
cleaning of crown, dying of basal, extremely shaded branches). Temperature
demands for the photosynthesis are rather lower than for respiration. The optimal
temperature of central European tree species is about 25 °C. Conifer needles may
have active primary photosynthetic processes even in temperatures as low as 0 °C.
In temperatures below zero, the photosystem II finishes.
High portions of solar radiation (long sunny days, around the noon) may lead to
the fusion of tetrapyrol from phytol (dephytolation of chlorophyll), which makes
chlorophyll inactive – the damaged part of a leaf is bleached. The acid
environment causes a replacement of a magnesium atom in a chlorophyll
molecule by a hydrogen atom. Moreover, this makes the chlorophyll inactive in

75
photosynthesis, its green colour changes into yellow-brown or brown (the affect of
acid rains, browning of fallen leaves during the decomposition).
The most common reason for limitation of photosynthesis is a lack of water, reduced
transpiration resulting in increased leaf heating. Besides inadequate transpiration
flow, photosynthesis could be limited by suboptimal mineral nutrition, namely the
nutrients necessary for this process – e.g. N, P, Fe, Mg, Cu, K, Mn, Cl. Moreover, a
lack of water in leaves brings about a closing of stomata and an increased
resistance of leaf epidermis to CO2 entry into a leaf. If water stress lasts longer,
photosynthates are not taken away from leaves but remain there and accumulate,
which substantially limits photosynthesis. The speed of photosynthesis is given by
the speed of gas diffusion (CO2, O2, water gases), which is driven not only by
external factors, but mainly by factors such as the position of a plant as a part of a
stand or as a solitary tree, the shape and arrangement of a crown and leaves, the
surface of leaves, the functions of stomata, the arrangement of mesophyll, the
system of conductive tissues etc.
Cells in green plants also contain other systems of molecules capable of utilization of
solar energy. However, their activity is more complex than in chlorophylls. Such is
phytochrome – a system of two forms of one molecule type made of a chromophore
and protein parts. P660 and P730 forms blend into each other after the absorption of a
photon of a corresponding wavelength and they make a photoreceiptor of plant cells
in the red light spectrum. Through phytochrome, light controls activity of a range of
enzymes, synthesis of nucleic acids, proteins, chlorophylls and other elements, but
also seed germination, flowering, leaf development and fall off, formation of buds etc.
8.1.2 CO2 exchange in plants

The cellular metabolism of carbon is connected with the outer environment via the
exchange of gases. Chloroplasts consume needed CO2 in photosynthesis, and
release oxygen. At the same time, cells constantly absorb oxygen necessary for
respiration and release CO2. In a given time any of the two contrary processes may
dominate in photosynthesising leaves. Respiration, active in the light, consists of
photorespiration and mitochondrial respiration. During the day, the speed of CO2
reception needed for photosynthesis related to a unit of biomass usually
exceeds the speed of CO2 release by respiration in the light. The result is a net
receipt of CO2 by a leaf. If the rate of photosynthesis falls down, it may happen that
respiration is high enough to balance it (it reaches the compensation point, see
above). If the rate of the photosynthetic activity continues to slow down,
respiration prevails, and in the dark, there is only a respiration release of CO2.
For instance, metabolism in shaded leaves in the crown base or inner periphery or a
crown core reaches the compensation point. It is a leafless space with inadequate
conditions for a long-term positive balance of carbon. The compensation point may
be even reached due to an increased temperature of leaves (above 35 °C), which
speeds up foliage respiration and slows down photosynthesis. A range of other
stressing factors including protective reactions after injuries etc may have a similar
effect.

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8.1.3 Photosynthesis and respiration

Photosynthesis and respiration are complementary processes inside plant cells,

i.e. leaf mesophyll cells where both processes happen simultaneously in
separated compartments of chloroplasts and mitochondria. On another level,
these two processes within one individual plant take place in green parts (leaves,
petioles, young stems etc.), while respiration only takes place in living cells with a
lack of light (cells inside branches, stems, trunks, roots). This means that e.g.
respiration of cells in the cambial zone also gives carbon dioxide for the
photosynthesis of cells in the primary bark. At a larger scale, we may consider the
relation between photosynthesis and respiration as similar to relations between
producers (green plants), reducers and consumers (heterotrophic organisms), at
first sight completely distinct from plants. This means that not all exchanges between
plants and animals are one-way. Plants, animals and bacteria on Earth have
coexisted for so long that some of these organisms have become a natural part
of the environment or even body of other organisms. Some CO2 molecules, now
fixed by the photosynthesis into organic molecules of a green leaf, could be released
into the atmosphere by breathing of an animal or by bacteria decomposing dead
organic matter a day before. So it is clear that the utilization of carbon makes a
cycle that involves the biosphere (all living organisms on Earth) as a whole and
extends boundaries among individual organisms. It is similar with atoms of
nitrogen, phosphor and sulphur moving between organic and inorganic realm, plants,
animals, fungi and bacteria.

8.2 Respiration

Cells gain energy via the oxidation of organic molecules. Earth atmosphere
contains a large percentage of oxygen. The bond with oxygen makes the most stable
form of carbon in CO2, and the most stable form of hydrogen in H2O. This means that
a cell is able to gain energy from saccharides or other organic molecules by allowing
their carbon and hydrogen atoms to bond with oxygen and the formation of CO2
and H2O. Then a cell releases this energy into macroergic bonds of ATP. This
process is called respiration and gaining energetic reserve of oxidative
phosphorylation.
All organisms, to be specific all their living cells, respirate. (Another mechanism
of organic matter decomposition is a quick oxidation in burning, e.g. wood, petrol,
natural gas)
The summary chemical equation of respiration is sometimes simplified and
expressed as the opposite to photosynthesis, i.e. as glucose oxidation and release of
energy, carbon dioxide and water:
C6H12O6 + 6 O2  6 CO2 + 6 H2O + E.

Yet, respiration is much more complex, it processes in many controlled gradual

steps and its outcome is a wide range of synthetic pathways and coordinations. That
is why we recommend that respiration is studied as a metabolic centre. Plant
respiration disintegrates complex substances with high potential energy (e.g.
saccharides, fats, proteins) into simple substances and a release of energy. This
energy serves for various cell functions – intracellular movement, maintaining
and renewal of structures, syntheses, release of heat (e.g. in spring in opening
buds and flowers). The disintegration of respired substrate (e.g. glucose) is a

77
several step process: glycolysis, pyruvate decarboxylation, acid citric cycle,
oxidation of final acceptor of electrons (NADH2). Glycolysis is localized in
cytoplasm, while acid citric cycle and oxidative phosphorization in mitochondria. The
yield of the whole disintegration is 36 mols of ATP (adenosintriphosphate), 2 mols of
GTP (glyceraldehyde 3 phosphate) and free enthalpy ΔG= –2,87 MJ per 1 mol of
glucoses. Besides the release of energy, disintegrative steps form a high number of
useful products that become preconditions for photosynthesis (e.g. lignin).
Respiration is more effective in presence of oxygen (aerobic respiration), but it may
even occur without a molecular O2, by means of a chain of reduction-oxidation
reactions forming simpler molecules, but still rich in energy. This is anaerobic
respiration, further classified into fermentation (disintegration of saccharides and
lipids) and decomposition (disintegration of nitrogen organic substances).
Fermentation is classified by the final, accumulated and unprocessed product e.g.
ethanol fermentation, milk fermentation, butter fermentation etc. These processes
predominantly give energy to microorganisms (bacteria, yeast etc.).
It may happen even to woody plants that conditions for aerobic respiration of
roots are not favourable due to flooding, compaction of soil. What is more, in
decreased soil aeration, anaerobic soil bacteria produce potentially toxic compounds
such as gases, organic compounds – alcohols, carbonyl group, volatile fatty acids,
non-volatile acids, phenolic acids and volatile compounds of sulphur. In such a
situation, even the development of tree shoots is affected due to elongation of
their internodes and limitation of formation and spatial growth of leaves,
induction of leaf aging, damage and fall off. In long term affecting, diameter
growth of adult trees slows down and the structure of phloem and xylem is
modified. Moreover, the formation and growth of roots is severely impacted, and
the root diminishment is intensified by acting of r. Phytophtora mildews. The lowered
respiration of roots results in limited photosynthesis rate since it induces the closing
of stomata and modifies carboxyl enzymes, and decreases the content of
chlorophyll. Absorbing of mineral nutrients falls with a lowered area of the root
system and with a lack of energy: energy release in anaerobic respiration is weaker
and does not meet the needs of root functions (ions from roots could be released into
soil). Modification and reduction of root growth make trees prone to windthrow. The
levels or auxine, ethylen and abscis acid in stem rise, while levels of gibberellins and
cytokinins fall. Globally weakened woody plants incline to attack of biotic
factors.
Plants, or their living cells, particularly meristems, respire continuously during
the day and night, photosynthesize only in daytime (light). Photosynthesis
prevails respiration in leaves several fold (approx. 5–10 fold). Respiration utilizes
energy most efficiently from simple saccharides that cannot be accumulated in
organisms in higher concentration since they are osmotically active. They typically
modify into other forms or reserve substances – polysaccharides (most frequently
starch), lipids (mainly as a reserve in seeds e.g. Juglans, Fagus, Corylus), and
sometimes reserve proteins. Structure elements of cell walls in woody plants cannot
be taken back as a source of energy even though they are saccharide type, such as
cellulose, hemicelluloses, pectates etc.
Finally, proteins, too, make the aerobic substrate in which aminoacids as foundation
stones could be reused for new or different, better functioning structures (e.g. in
reconstruction of cell walls).

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8.2.1 The process of respiration – oxidation of glucose

Respiration is a two step process – it begins by anaerobic glycolysis and continues

by aerobic disintegration of pyruvic acid or fermentation.

8.2.1.1 Anaerobic glycolysis

It takes place in cytoplasm along with catalytic activity of enzymes dissolved there. It
is only active without the presence of oxygen, even in cells located in aerobic
surroundings. These two facts prove the old origin of this metabolic pathway. The
final product is two molecules of pyruvic acid and ATP.
8.2.1.2 Aerobic disintegration of pyruvic acid (in mitochondria)

This process happens into these three stages: 1) decarboxylation of pyruvic acid,
2) citric acid cycle and 3) respiratory chain (Fig. 38):
1) decarboxylation of pyruvic acid:
One atom of carbon in the form CO2 splits off from pyruvic acid (CH3COCOOH),
which means the acid is decarboxyled, and in co-working with acetyl coenzyme A,
activated acetic acid is formed
2) citric acid cycle (Krebs cycle):
It progresses in a matrix. The activated acetic acid reacts with oxaloacetate,
which results in the formation of citric acid. A molecule of ATP and hydrogen are
released. Three molecules of coenzyme NADP+ and one molecule of coenzyme
FAD (flavin amide dinucleotide) reduce by accepting hydrogen and take it in the
respiration cycle.
3) respiratory chain:
This is a sequence of oxidation-reduction reactions. Protons as well as electrons
bring reduced coenzymes NADPH + H+ and FADH2. At the beginning, protons get
released into the neighbourhood. Electrons are transmitted by a range of transmitters
into the inner membrane of mitochondria in the gradient of electric potential, and
ATP is synthesised at the same time. ATP synthesis in the respiratory chain is
called oxidative phosphorylation. It is very similar to phosphorylation taking place
on thykaloids in photosynthesis. Transmission of electrons generates energy
which allows for the transmission of protons via the membrane from matrix to
intermembrane space to get accumulated. This is a process of foundation of
mitochondrial membrane proton gradient (gradient pH) induced by different
concentration of protons in the matrix and intermembrane space. Protons get back
into the matrix through an ATP enzyme that puts energy released during
transmission into ATP. Transmission of two protons into matrix is necessary for the
synthesis of one ATP molecule. More ATP is generated in phosphorylation than in
photosynthesis.

At the end of the chain, electrons are transmitted to the molecular oxygen. This forms
oxygen anions, that later join with protons to make water

79
2 H+ + O2– = H2O
In oxidation processes in mitochondria, 36 molecules of ATP per one molecule of
glucose are generated.
Enzymes catalysing reactions of Krebs cycle and respiration chain are located and
originate in the inner membrane of mitochondria.

Fig. 38 Schema of main metabolic processes in plant. Products of photosynthesis are

directly deposit to storage reserve (starch), cell wall (cellulose) or they entry to five
carbon-sugar cycle, to nucleotides structure. Saccharides, fats and proteins can entry
to process of respiration. Not only energy is produce during respiration, but also other
substance are formed, which are the basis for the secondary plant metabolisms
(polosaccharides, lipids, proteins, phenols, isoprenoids, hormonones and many
others). Thus, respiration has the key position in the plant metabolisms.

8.2.1.3 Fermentation

Pyruvic acid is degraded without the presence of oxygen and reactions stop
when a semi-product is generated, e.g. ethanol. This is a case of ethanol
fermentation occurring even in plants in anaerobic conditions (see above). Ethanol
80
is toxic, particularly to the activities of mitochondria; longer anaerobic conditions
(mainly in summer) may lead to dying of plants. More resistant woody species
include the ones that occupy meadows with regular spring flooding or the trees living
in higher altitudes near mountain streams – quick and cold water is rich in oxygen.
Less energy is generated through fermentation than through aerobic respiration
as most energy is bound to final outcomes.

8.2.2 Factors of respiration

In individual plants, flowers and unripe fruits respirate faster than leaves, roots
respire faster that stems of shoots. The main areas of respiration in branches and
trunks are in bark – mainly phellogen, living part of phloem, cambial zone and
outer functional layers of xylem (sapwood) containing living cells. Growth
respiration (breathing in growing season during growth and synthetic processes in
cells) in seedlings, root and stem apices, in opening of leaves and development of
fruits is from 3 to 10 times more intensive that the maintenance respiration (e.g.
winter breathing of needles). The rate of respiration is directly proportional to the
rate of growing. As differentiation and maturation of tissues proceed, respiration
activity weakens. A temporary, sharp rise in respiration occurs during fruit
ripening and leaf maturation (i.e. climacteric respiration). This is a breakdown of
substances rich in energy, withdrawal of mobile mineral nutrients (e.g. potassium,
nitrogen), together with an assimilation of toxins from metabolism into a structure that
is to be separated from a plant and for a preparation of the separation, i.e. healing
layer. These enzymatic processes of natural maturation are under a strict genetic
control and depend on favourable conditions of the environment. They are reflected
e.g. in a gradual change of leaf colour in autumn (cells surrounding venation live
longest). It is not the same as a sudden dying (e.g. due to drought), in green state
and a postponed fall off. Dying of plant cells is often accompanied with a release
of gaseous metabolic products (e.g. ethylene).
In winter, cells in ash branches work only to survive and respiration is limited to
approx. 2 mg CO2 per hour per 100 g of fresh biomass. As soon as the growth
finishes in September, respiration weakens. In woody plants, respiration increases
during the secondary diameter growth and remains very active till late August, in
some species even longer, e.g. oak and beech till late October, lime as late as
December. Respiration in winter is balanced. It is one third to one fourth lower than in
high summer. So the rate of respiration does not only alternate during the day but
also during the year. Different plant species demonstrate different rate of
respiration in the same conditions. For example, the average speed of breathing
in leaves of deciduous trees is five times higher than breathing in assimilation organs
of evergreens. The rate of released CO2 and accepted oxygen (respiratory quotient)
is typically one. This happens only if starch is burnt in respiration, but if fat is used
(fatty seeds – Fagus, Juglans), the respiratory quotient falls bellow one. Yet, from the
perspective of a plant sustainability, the total yearly balance between
photosynthesis and respiration is required to be positive, i.e. photosynthesis
needs to dominate over respiration. Otherwise, growth could be endangered.
The rate of respiration is affected by a number of endogenous and exogenous
factors. The endogenous factors include (excluding species and individual
exceptions):
1. growth

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 2. available substrate, i.e. concentration of organic compounds in cells (the
higher the concentration, the higher the rate of respiration)
3. amount of water in tissues.

Seeds begin breathing more intensely as soon as they soak with water. The
exogenous factors mainly include:
1. temperature: respiration intensifies in higher temperatures (to 40 °C), and
weakens in lower temperatures. Pine needles respire 25 times faster at 0 °C that
at –12 °C. In the temperature range 0–40 °C, the speed of respiration grows
twice to three times every plus 10 °C, but between 50 °C and 60 °C enzymes and
functionally significant membrane structures are damaged by heat and
respiration dies down;
2. light does not affect respiration;
3. air humidity is a positive factor;
4. a higher CO2 concentration in the atmosphere slows down respiration,
40 % concentration is harmful;
5. injuries enhance respiration intensity (related to an active preparation and cell
division).

As little as 1–3% concentration of oxygen will be sufficient to a normal progress

of mitochondrial respiration. If concentration falls bellow this limit, at first
respiration slows down, and then the intake of water, ions and root growth decrease.
Abscisic acid production rises, especially in ethylene, so the above ground growth is
interrupted and leaves fall off prematurely. Inadequate conditions may launch
respiration in higher plants without oxygen (anaerobic respiration, see above). This
may happen when roots are flooded or when a seed germinates in the muddy soil.
However, this is only an extemporary solution. If plants lack oxygen for longer, they
die. Plants on extremely heavy soils with a slow movement of oxygen develop an
extensive root system and adventitious roots. Pneumatophores (aerating roots)
may be developed as an extreme specialisation. Plants usually use saccharides for
respiration. If they happen to require some other substances, such as lipids, they
modify them into saccharides by themselves. It is only an exception that a plant uses
proteins for respiration.

8.3 Photorespiration

A metabolic process similar to respiration is active in plant part which contain

chlorophyll. This process co-works with photosynthesis, in which O2 is used up
and CO2 is released in the light, but, in contrast to respiration, this is not active
in the dark. This exchange of O2 for CO2 is called photorespiration. It begins in
chloroplasts, moves to peroxisomes and finishes in mitochondria. The enzyme
of photorespiratory metabolism is enzyme RuBP (Ribulose-1,5-bisphosphate
carboxylase oxygenase). During the intake of oxygen this enzyme splits ribulose-
1,5-bisphosphate which releases CO2, having been taken in before. The rate
between photorespiration and photosynthesis is controlled by the input of O2 and
CO2, radiation density, temperature etc. A high partial pressure of O2 stimulates
photorespiration, while photosynthesis is stimulated by a high input of CO2.
In average conditions (21 % O2 and 0,03 % CO2 in the atmosphere, intensive
radiation, temperatures between 20–30 °C) plants lose about 20%, extremely up
to 50 % of CO2 gained through photosynthesis in the form of photorespiration CO2.

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On the other hand, this process uses energy (prevents from leaf overheating) and
provides precursors for the renewal of chlorophyll.

8.4 Water regime of woody plants

Functions of all living organisms on Earth directly depend on characteristics of

water and substances that are either dissolvent in water (ions of inorganic salts,
monosaccharides, proteins, organic acids etc.), or wettable (hydrophilic),
hygroscopic and soaking (cellulose, wood, colloids), or, on the other hand, are
water repellent-hydrophobic (fatty substances, lipids).
The most significant physical characteristics of water include:
- specific heat of ice melting (amount of energy needed for the transition of a
unit of mass from solid state to liquid state) is 334 kJ.kg–1,
- specific thermal capacity (amount of energy needed for heating a unit of mass
of 1 °C) of water in liquid state is 4 186 kJ. kg–1.K–1,
- specific boiling heat (amount of energy needed for the transition of a unit of
mass from liquid state to gaseous state) is 2 260 kJ.kg–1,
which are extremely high values in comparison to other substances. In transition from
the liquid to solid state there is an increase in volume, meaning a reduction in density
(i.e. ice floats on water). Water density is almost exactly 1000 kg.m–3, i.e. one litre of
water weights one kg.
Volume heat expansion is negligible in both the states (liquid and solid) compared to
volume changes during melting or freezing. We could say that water rarely changes
its volume with changing temperatures, which is true for both the stages – liquid or
solid. Nonetheless, there exists a little relation to temperatures, as it is measured that
the highest density of water is at the temperature of 4 °C.
Like all liquids, water has a measurable surface tension. This is energy needed to
extend a liquid surface area of 1m2. This value equals 0.072 J.m–2 (e.g. water in
weightless state has a spherical shape, with the smallest area related to volume).
Viscosity tells us how easily a liquid can flow. The lower the viscosity (surface
tension), the weaker the resistance of a liquid e.g. flowing through a pipe (through
vessels in case of woods).
The electrical conductivity of pure water in normal conditions is so low that it could be
considered a great non-conductor. A negligible electrical conductivity is caused by a
little amount of dissociated molecules (present in the form or hydronium ions H3O+
and hydroxide ions OH–). Nonetheless, dissolvation of all other elements that tend to
dissociate may increase the conductivity of water solution to several fold.
A molecule of water contains two hydrogen atoms and a central oxygen atom that
form an angle of 105° (in solid state – ice); this angle slightly oscillates in a liquid
state. Even though the water molecule behaves neutrally, protons positioned in this
angle from the oxygen atom surface cause a certain polarity, i.e. a weak positive
charge on one side of the molecule. It results in an attraction of the positive charge
side of a water molecule by a negative side of a neighbouring water or other
molecule. These weak interactions appearing in a range of organic molecules
(energy from 8 to 42 kJ.mol–1) are called hydrogen bonds. In case of water, these
are the factors responsible for its abnormal performance – being liquid in a range
from 0 to 100 °C, being a polar dissolvent, forming true and colloid solutions. The
cohesiveness of water molecules among each other is called cohesion, the
cohesiveness of water molecules with other molecules is called adhesion. Cohesion

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and adhesion of water together with hydrogen bonds with a range of other structures
secure compactness of plant bodies.
A constant volume and non-compresiveness of liquid water is responsible for the
diameter growth of cells and the self-supporting capacity of unlignified parts of
plants (leaves, young stems etc.). Water inside cell vacuoles develops an internal
pressure on the cell wall (turgor). In balanced pressure of vacuole and resistance
from the cell wall, there is a state of fulfilled cell turgor. If cells are turgescent, they
develop a mutual pressure and tissues remain stable. In loss of water, they wither.
There are numerous cases when cell walls in tissues are arranged in such a way that
they attract molecules of water in various ways and various directions. A change in
water content leads to movements of the whole organs or their parts – e.g.
change in leaf position in space and tilting, quick movements of leaves after
being touched (Mimosa), opening and closing of pores, movements of flower
sections, opening and closing of cones and changes in a position of the whole
branches as a result of presence of reaction wood.
Values of specific heat of ice melting, water boiling and specific heat capacity of
water tell us a lot about the exceptional performance of water in plant tissue
thermoregulation. Parts of woody plants with a high content of water (apices, fleshy
fruits of woody species, cambial zone etc.) have a high thermal stability. The
consumption of heat in water vapour from leaves (transpiration) is very important for
cooling of these. On the contrary, in the process of water freezing, cooling of tissues
slows down. The described water volume changes during melting and freezing are
a primary factor of damage of plant tissues, particularly in repetitive or quick
fluctuations of temperatures around 0 °C . Other water capabilities such as
performing as a reagent, donor and acceptor of electrons and as a mobile
dissolvent point out to its extraordinary significance that cannot be substituted.
Gradients in rainfall, contrast expositions and differences in texture and depth
of soil, that play a vital part in water balance of plants, have an impact on
vegetation, its types and functional state. Water also plays an important role in
decomposition, erosion and control over nutrients in the soil. Water regime is
closely tied to mineral nutrition, photosynthesis, respiration, growth and
architecture of plant organs.

8.4.1 Water content and its thermodynamic state

From the perspective of phylogenetics, life originated in water, and water remains
the primary medium of biochemical processes in plants. Cytoplasma contains almost
90 % of water on average (of fresh mass weight), organelles rich in lipids as well as
chloroplasts and mitochondria contain around 50 % of water, and, regarding the
whole organs or tissues, leaves and fine roots contain about 80 %, conductive
parts of xylem (sapwood) about 50 %. Above-ground parts of terrestrial plants
permanently lose water by evapotranspiration. By the ability of plants to
compensate short-time oscillation of water content and to sustain a loss of water,
these types of plants are distinguished: poikilohydric and homoiohydric.
Poikilohydric plants (fungi, some algae and lichens) adjust their water content to the
humidity of the neighbourhood – they contain small cells without central vacuoles. If
dry, their volume diminishes and their vital functions are suppressed without any
damage to protoplast. Cells of homoiohydric plants contain a set of vacuoles that
can balance the impact of extragenous humidity conditions to a certain extent. The
development of a cuticle, trichomes, control of transpiration through pores,

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suberinized surfaces and ventilation systems, the development of long distance
conductive systems and complex root systems securing water uptake, all of these
have made terrestrial plants capable of retaining their inner balance even in
sudden changes in humidity in their neighbourhood, and this capability has
allowed for the global spread of plants. Even homoiohydric plants have stages when
they perform as poikilohydric and can sustain significant losses of water. This
concerns ripe seeds that retain the germination capacity for a long time, up to
hundreds of years by minimisation of vital processes along with a radical limitation of
hydratation (water content may fall below 10 %).
Most of hydratation water is bound in cytoplasm and cell wall, in the power of 1–10
(or more) MPa, depending on the density of cellulose fibres. Forces that bond water
to structural elements of a matrix (cell wall, colloids) are defined as matrix potential.
In the qualitative point of view, the significance of water contained in a cell grows with
the rate of its availability. The most accessible water is present in all vacuoles,
i.e. cell compartments specialized as water reservoirs, or in dead cells serving as
hydrocytes. Over a half of water content in leaves is accumulated like this.
Though, neither this water is completely mobile, as it is osmotically bound to
dissolved substances such as saccharides, organic acids, secondary metabolites of
plants and ions. Forces that bind water in dilutions are altogether called an osmotic
potential. Only purified water has the highest specific free energy, i.e. potential
to perform activities (e.g. dissolve ions, cover coloid particles...) and to move
freely. Water potential of chemically pure water is a foundation for expressing a
relative chemical potential of water which is tied by various forces in any place in the
system of soil-plant-atmosphere. The chemically pure water potential is the highest,
meaning zero, and the potential of water bound in systems lowers, e.g. is
negative. It depends on the above mentioned osmotic potential (concentration of
osmotically active substances, oligosaccharides in plant, salts in soil), pressure
potential (turgor stress on cell walls from the inside of cells and hydrostatic pressure)
and matrix potential (adhesion to cell walls, water cohesion of colloids etc.). Water
potential may be understood as an activity needed to develop to increase the
potential of bound water up to the grade of pure water. It is usually expressed as
an amount of energy per volume unit (J per m–3), as a unit of energy per amount of
material (J mol–1) or as pressure (MPa).
The osmotic potential of a solution falls down not only with the growing number
of diluted substances but also with the rising temperatures. Macromolecular
substances may be present in high weight amounts without any significant decrease
in osmotic potential. Though polymeration of small molecules to macromolecules
(e.g. change of saccharide into starch and this process reversion – hydrolysis) may
highly affect osmotic potential of a solution, which means that the net flow of water
may be controlled by plant. In protoplasts with a central vacuole, there is a close
relation between osmotically bound water in a cell and its availability in cytoplasm.
Water availability is expressed as a total water potential of the water system
(e.g. cells, cell compartment or outer solution, usually symbolized by Ψ), which
means that the more negative the water potential of a given system, the lower the
water availability from the system to the neighbourhood. An osmotic component of
water potential depends on the character and concentration of a dilution and is
expressed by Van Hoff equation

Ψπ = –Φ R T ρw C = Φ R T (Ns/V)

85
R – universal gas invariable (8.31 mol–1 K–1)
T – absolute temperature (K)
ρw – water density
C = Ns/V is a sum of moles in a dilution per a unit of water weight in symplast, or
water volume in a cell V
Φ – osmotic coefficient that includes non-ideal characteristics of a given dilution in
the thermodynamic perspective.

Besides the osmotic constituent of water potential, the total water potential of
water system contains matrix and pressure potential constituents.
A difference in potentials between places with different values of water potential (Ψ)
is a situation analogical to electrical circuit with points of different voltages (U)
measured in Volts. Electrical current tends to flow from a place of a higher voltage
into a place of a lower voltage in the circuit. It is similar in a living plant; the current of
water flows from places of higher water potential (positive, i.e. less negative)
into the places of lower water potential (more negative).

8.4.2 The movement of water in a cell

There are two mutually interconnected water circulation systems in woody plants.
Water circulation in a cell (small circulation) and circulation in the whole plant
(large circulation). The small circulation secures a continual exchange of substances
in a cell, and the large circulation secures transpiration and assimilation flow.
All the processes in a plant may only be realized in the state of sufficient
hydratation. Water moves from places of a higher water potential (average values
in soil –0.1 to –0.25 MPa) into places with a lower water potential (e.g. to roots with
values –0.2 to –0.3 as far as leaves with a lowered water potential –2 to –3 MPa, and
then into the atmosphere with values of dozens or hundreds of MPa). Another way of
balancing different gradients of water potential is a direct movement of diluted
substances (diffusion, occurring only in the apoplast, e.g. in intercellular spaces).
Though, this is limited by semipermeability of vacuole membranes (tonoplasts)
and cytoplasm (plasmalema), that only free water molecules. The output process is
osmosis. Water flows into a cell situated in the environment of a higher osmotic
potential (hypotonic) from another place if a cell wall resists the pressure from the
inside. If a cell wall solidity is lower, a cell bursts (plasmoptysis). On the other hand,
in conditions of a lower osmotic potential, water flows out of the cell and protoplast
limits its volume (plasmolysis). If a plasmolysed cell is transported into a hypotonic
environment, it may be returned into its former state by means of deplasmolysis.
Otherwise it dies. Plasmolytic processes are important for the diagnosis of a health
state of cells and cell membranes. If they are damaged or injured, substances from
its neighbourhood are free to enter the cell, and material from the inside of a cell can
move out. Protoplast loses its capability of selecting passing substances.
If cells are only partially saturated with water, there is a state of hydratation deficit.
This is a cause of a water flow in a cell, or in a plant. If water deficit exceeds a certain
limit, life sustaining processes in a cell weaken.
A plant needs different amounts of water in different life stages and a water
potential of cells changes by this.

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8.4.3 Water saturation deficit

Water saturation deficit (WSD) is an amount of water that a plant lacks to its full
saturation. It is expressed in percentage and related to the maximal content of water
in a plant. Variables of the original weight, weight after saturation and weight of a dry
mass are studied in order to calculate the deficit.
After a little loss of water and its regaining, plants renew their natural colour and
turgor of cells. Such a water deficit that will suffice for re-saturation of cells without
any damage is called a critical water deficit. A sublethal deficit is such a state that
projects in a plant the first damage and a lethal deficit is a state of a serious damage
to a plant, disabling a full saturation in the future. Water deficit has a grave impact
on metabolism and physiology of cells, it slows down primary life functions of
a plant and its growth.

8.4.4 Water potential of plants

Water potential of a plant is related with other physiological processes and its annual
growth. An inner constituent of water potential is cellular water potential;
exogenous factors include availability of physiologically present water, solar
radiation, temperature and variations of temperature and other factors.
Values of water potential of plants change during the day. Minimal values occur in
the midday or in the early afternoon. Due to irrigation, the value of water potential
of root tissues quickly rises.

8.4.4. 1 Water potential of soil

Soil humidity is the total content of water in the soil at a given moment and in
given conditions. The mobility of soil water and forces that tie water in the soil
are factors that influence usability of soil water by plants. The total amount of water in
soil is called the water capacity of soil. It mainly depends on the size of soil
elements. The smaller the average size of soil elements, the larger the water
capacity of soil. A plant cannot utilize all water contained in the soil. Water
potential of soil is the decisive factor of its usability by plants. It is a vector sum of
forces that bind water in the soil. It comprises of the total water content in soil and
osmotic value of soil solution.
If we take into consideration various forms of water in the soil – hygroscopic,
adhesive, capillary and gravitation water, only capillary water is physiologically
utilizable to plants. Differences between the absolute and biologically utilizable water
content in different soils and substrates are significant for the determination of a
wilting point. This point is a critical content of water in the soil below which a plant
starts withering.

8.4.6 The uptake and conduct of water

8.4.6.1 Transport of water from soil to roots

Water passes into a root through rhisodermis. Then it migrates through cell walls
of the primary cork to cells of xylem parenchyma. The movement of water from soil to
root is realized by forces of apoplastic movement allowed by cell wall micellae and
a colloid state of cell content. The following movement of water in a root is realized in

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a symplastic way through cytoplasm, except from vacuoles, and through
plasmodems. The last water movement is driven by osmotic potential among
vacuoles. Water potential falls from the surface towards the central cylinder. A sharp
fall in a water potential occurs in the transition of cells of storage parenchyma through
endodermis into a xylem conductive part. On one hand, cells soak water in, and on
the other, they push it out actively.
Differences in the values of water potential in individual parts of a cell (water potential
in the outer side of a cell is higher than in the inner nearer the root centre) are
conditioned by the activities of a living protoplast. Water transport is not realized
only in one way, but it a complex process.

8.4.6.2 Water conduction in a root

There are two mechanisms of water uptake by the root system: active and
passive. The active uptake is characteristic for a low release of water by a plant,
the passive intake occurs in an intensive release of water.
The active uptake (biological) of water by the root utilizes energy of material
metabolism. The loss of water in transpiration brings about an under pressure, which
leads to soaking of water in the passive uptake.
The uptake and movement of water in a plant is well described by the process of
electroosmosis, meaning the movement of a dispergent phase (water) in an electric
field towards an electrode with the opposite charge. The electroosmosis is only active
if a disperged phase is immobile, and if only the disperging environment is
mobile. In plant cells, elements of the surface layer of immobile protoplast are ether
positively or negatively charged, which means that a water solution moves towards
the negatively charged inner surface of protoplast. The outer and inner layers of
protoplast form a boundary between the water solution (as the outer environment),
vacuole and cell flesh (as the inner environment), so these are media of electric
charges. There is a potential difference between these two bordering surfaces. Water
solution contains ions of opposite charges and hydratation coat, which carry water
adhering to them; the lower the solution concentration, the more water passes
into cells. Yet, electroosmotic passing of water into cells greatly depends on the
intensity of life processes (respiration) and on conditions of the exogenous
environment. Due to the described processes, there are changes to the electric
potential on the surface of protoplast.

8.4.6.3 Root pressure

Movement of water through a root develops a pressure that pushes water upwards
into higher tissues of a plant. It is caused by forces of colloid systems and osmotic
forces of protoplast or root cells. Root pressure is a physiological process requiring a
high amount of energy. It is active in spring by means of an intense hydrolysis of
starch into dissolvent saccharides, e.g. in tree roots. This process limits a osmotic
constituent of water potential of a root structure. When the soil has suitable
humidity, soil water flows into roots due to electric gradient, and is further
pushed upwards into the above ground system. After watering, budding and
development of leaves, the root pressure falls down.
Root pressure may be demonstrated during plant injuries in spring prior to budding of
leaves. A liquid flows off a scar after a trauma (e.g. Betula, Vitis vinifera etc.).
Besides water, this liquid contains organic substances such as saccharides, amino

88
acids, organic acids, enzymes, phytohormones, mineral salts etc. This "bleeding" is
economically utilized for gaining organic material in many industrial branches (e.g.
sugar from Acer saccharum). Root pressure is highest in the morning and in the
evening, which corresponds to the values of water saturation deficit and pressure
potential. Root pressure only develops a slow movement of water and is not able to
transport large amounts of water quickly into the higher locations.

8.4.6.4 The impact of various factors on root uptake of water

Environmental factors or water uptake by plants are direct or indirect. Direct forces
on water uptake by roots are the ones that control transpiration of plants (release of
water). Other ones include factors that control root growth, since water is only taken
in by young growing roots. Soil conditions belong to indirect factors, for example
the content of water in soil, temperature and aeration of soil, water potential of soil
water values etc.
The soil temperature considerably affects water absorption by roots. If lowered to
3–5 °C, the above ground parts of a plant wither even if they give out only little water.
On the other hand, as soon as the temperature rises to 12–15 °C, plants return to
their normal state. Thermophilic plants limit water uptake at temperatures as high as
10 °C, and they completely stop this activity at 2–4 °C. On the contrary, plants
adapted to cold temperatures (e.g. ash, birch) take in water even at lower
temperatures about 0 °C. The optimal temperature for intake of water is about 20 °C
for the majority of woody plants. The best conditions for a fluent intake of water are
when soil temperature is 2–5 °C lower than air temperature. This corresponds to
the temperature gradient of different species. Though, cold soil is physiologically dry
no matter how much water it may contain (higher altitudes). Low temperatures
affect protoplast viscosity, which results in the changing in passing of water
through the membrane system of root cells. Temperatures, too, have an impact on
diffusion and other physical and chemical processes in root cells.
Soil aeration, mainly the rate of carbon dioxide and oxygen, controls water uptake
by the root system. Woody plants need oxygen for respiration, growth, metabolism
and the normal activity of the whole root system. A lack of oxygen limits respiration,
which further affects an active uptake of water and nutrients. The optimal content of
oxygen in soil is about 10–12 %. Too low or high content of carbon dioxide limits,
sometimes even stops, the uptake of water, as well. A rate between oxygen and
carbon dioxide that does not comply with optimal plant activities occurs on
compressed or flooded soils. The water uptake can only be satisfactory if a
sufficient amount of physiologically available water is present in the soil. The optimal
water uptake by plants occurs at 60–70% of maximal capillary water capacity of
soil. The most important is the soil layer with roots, making a physiologically active
profile (vegetation layer). This depends on a species and age of an individual and on
characteristics of soil layers.

Soil solutions need to have a really low concentration (0.02 to 0.05 %) in order to
sustain a fluent water uptake. Electrolytes enhance water permeability of a cell,
while non-electrolytes limit it. An experiment comparing solution concentrations
demonstrated that a solution of the –0.03 MPa osmotic potential enabled plants to
take in more water (up to 1.85 times more) than a solution of –0.18 MPa osmotic
potential in case they could not quickly adapt to the environment. In changing
concentrations of a soil solution, the water uptake more or less slows down in relation

89
with mineral nutrition factors. It is mainly the nitrate-nitrogen that can boost the
uptake of water by roots.

8.4.6.5 Transpiration flow

Water taken up by the root system (in exceptional cases a little percentage also by
the above ground structures) is conducted in various directions and to different
distances into all body parts. At first, water trespasses a root radially from
rhisodermis to xylem conductive pathways; this is a short or medium-distance
transport. The second stage includes a long distance transport of water via
xylem elements. In the third stage, water is transported from conductive pathways
(xylem) through leaf mesophyll to epidermis cells in leaves till it evaporates out
into the atmosphere.
Water taken up by roots sets out on a way that could be very long, often about 100 m
in trees. Water can successfully overcome long distances only via vascular
bundles. The driving force of the flow is a gradient in water potentials between
two opposite apices. The flow could be broken by a lot of resistance factors such
as frictional force, gravitation potential etc. Plants, especially woody species,
have developed a perfect systems of tracheids, vessels or both for the conduct of
water on long distances. Water moving upwards via tracheids is filtered through
pit pairs of their tilting closings end walls. Tracheids are narrow, their
conductiveness is slower than in the case of vessels. Vessels consist of individual
vessels elements standing one on top of another, their cross compartments are
performed to a different measure. The weakest resistance of water conduct is
developed by liana vessels. Movement of water in deciduous woody plants is
easiest in the vertical direction. It is slowest in conifers since the pathways are not
vertical, but they are often spiral. Neither is it easy for water to move in the tangential
direction into phloem-xylem rays that distribute water in the radial direction with a
slight gradient of osmotic potential.
The highest stage of evolution is demonstrated by the conductive tissues in trees.
The youngest annual rings and young wood conduct water best, while the
central wood does not conduct water at all.
The movement of water in vessels is completely different from the movement in
parenchyma cells. The current of water in vessels is directed by the rules of
hydrodynamics while in living cells water moves by the rules of osmosis and
overcomes the resistance of cell walls.
The most important factor in water upward movement from roots into leaves is water
cohesion and adhesion to cell walls. Cohesion forces can lift water above
barometric pressure level and reach –30 to –35 MPa.
Water flow from root to transpiring parenchyma: parenchyma consists of cells with a
low water potential (several MPa, in woody plants –5.0 to –5.5 MPa). These cells
attract water from vessels. This generates a voltage in vessels between water film
molecules that move along the walls, so it appears as if all water was hanging on leaf
cells. If there is a sufficient amount of water in soil and if vapour is weak, root cells
support the water continuum in vessels, thus decreasing its voltage and enhancing its
upward flow. Besides water potential differences, another force of water
movement is the transpiration pull (i.e. energy of solar radiation), root pressure
to a certain extent (mainly in night irrigation), capillarity and changes in flow
sections of conductive pathways from roots towards leaves.

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8.4.6.6 The speed of water conduct

The speed of water conduct depends on a species, ontogenetic stage, on exterior

conditions, growing season (faster in spring). If physiological functions are in a
healthy state, the water movement in a plant is a continuous transpiration flow. The
speed of transpiration flow is expressed by an amount of water transported via
a certain part of conductive tissues in a unit of time and its fourth power
depends on the area of a cross section of a conductive pathway. In lower non-
vascular plants water only spreads by means of diffusion which reasons relatively
small growth of these plants. Water flow in vascular plants is faster. If the
transpiration flow is maximal, the average speed of water flow through xylem
of evergreen conifers is 1.2 m per hour–1, in deciduous scattered porose
species such as birch or lime it varies from 1 to 10 m per hour–1, and in an
instance of oak this range is between 4–44 m per hour–1 (circular porose
species), and up to 150 m per hour–1 in lianas.
8.4.7 Release of water by plants

Especially leaves, but also stems (mainly young ones) may be studied as bodies
saturated with water or humid areas that continuously release water in the form of
vapour into the air. The process of water evaporation from the surface of leaves
and other parts of plants into the atmosphere is called transpiration. It is driven by
physical rules of vapour and a physiological state of transpiring tissues. Plants leaves
can also release water in a liquid state by the process of gutation.

8.4.7.1 Guttation

A lot of plants are able to exclude water in a liquid state. This way of water release
is known as guttation (from Lat. gutta – drop). A plant presses water through special
pores – hydathodes. The cavity is filled in with thin-walled parenchyma cells, i.e.
epithem that encloses a vascular bundle bringing water. The root pressure is a co-
factor in guttation.
Guttation is not as frequent in woody plants as in herbs. It occurs when the air is
supersaturated with vapour, e.g. after a warm night or in early spring in the morning
when root pressure rises. Guttation is not the same as dropping of saccharide
solutions e.g. from lime leaves attacked by aphides.

8.4.7.2 Transpiration and its biological control

Transpiration is a loss of water from plants by means of evaporation (diffusion

process). It depends on physical conditions of vapour (humidity, temperature and
movement of air), water supply, plant species and ontogenetic stage, e.g.
rhythm. The rate of transpiration (amount of water released by a defined part of a
plant in a unit of time) varies within an individual (leaves in various positions,
radiation, age) by kinds of plant surfaces. Transpiration by stomata into the air is
called stomatal transpiration, transpiration by cuticle is called cuticular
transpiration, and a release through bark is known as peridermal, rhytidomal or
lenticelar transpiration (Fig. 39). Stomatal transpiration is easy to control for a
plant – closing of stomata rises a diffusion resistance of leaves up to 50 times. A
cuticular transpiration in which water overcomes the resistance of epidermis cells,
may reach up to 30% of the total transpiration in hygrophilous species, 3–10 % in

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conifers and only 1–2 % in succulents. The peridermal transpiration (from the
surfaces of suberionized stems) is comparable to the cuticular transpiration, in
summer it makes about 1 % of potential evaporation. The cuticular transpiration in
older well developed leaves presents about 1/10–1/20 of the stomatal transpiration.
The cuticular transpiration is affected by decreasing air humidity (e.g. fall of air
humidity from 95 % to 50 % brings elevates the cuticular transpiration about 5–
6times), rising temperatures and pollution of air that endangers wax surfaces of
leaves.
Water and nutrients are brought into living photosynthetising organs by means
of a transpiration flow, and it is again transpiration that cools these organs,
thus protecting them from overheating.
Woody plants that grow on dry sites have thicker cuticles, small numerous
stomata arranged in groups, able of quick reactions.
The rate of transpiration changes with age. It is more intense at first stages of
organogenesis than in older ones. Injuries of plants typically enhance
transpiration. Cut-off parts increase transpiration in the first minutes, yet, it later
slows down as a result of closing of pores.
Stomata make epidermis a partition with numerous openings. Water
transpiration through this partition is always higher than the transpiration from
an open water level, since molecules of vapour are not squeezed perpendicularly
above the surface. They form a fan, a shape supporting diffusion of the molecules
into the air. A lot of woody species face an excessive transpiration by enlarging the
border layer of air over a transpiring leaf surface and forming trichomes (Quercus
pubescens).
The area of stomata in leaf present from 0.6 to 1% of the total leaf area (Fig. 39).
The rate of transpiration through stomata is affected by the size, not by their number.
Opening of stomata immensely increases transpiration, but once opened,
transpiration slows down (Stefan rule of diameters). It is similar with closing of
stomata: rate of transpiration decrease is not linear.
There is a daily rhythm of opening stomata in the majority of woody plants: there
are two transpiration daily highwaves with a midday depression. In lack of water
stomata only open in the morning. The regulation is connected with a change in
turgor of stomata cells. Stomata are hydroactive and photoactive. Water deficit
affects stomata more than light. In lack of water, they even close in the light.
Permanent darkness elicits their opening, high density of radiation elicits closing, as
well as short-wave rays.

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Fig. 39 Stomata on the lower epidermis of oak leaf. Two guard cells can regulate
transpiration by increasing or decreasing the size of intercellular space between
them. Cuticular transpiration is lower by dense cutin layer.

8.4.7.3 Environmental factors of transpiration

8.4.7.3.1 Air humidity (vapour pressure in the atmosphere)

Air surrounding a plant may contain various percentages of vapour. Even air that
appears dry contains vapour. The absolute content of vapour is expressed either
by the partial pressure or by weight units. Nonetheless, the content of vapour in the
air could not exceed the level of maximal saturation (maximal pressure of vapour),
as being in a dynamic balance with a water level of the same temperature. It is
not usual for the air surrounding plants to be fully saturated with vapour, which
means it takes water in the phase of vapour from plants containing various
percentages of water. The movement of water molecules between a plant surface
and the air around a plant is expressed as a rate of vapour pressure on the plant
surface and in the surrounding air.
Air humidity is commonly expressed as a percentage relative air humidity (r) and is
calculated from the absolute humidity (e) and full (maximal) air saturation with vapour
(E), expressed in this equation:
e
r   100
E

The equivalent relative air humidity (rek) means a relative air humidity, yet it does
not calculate with air temperature (t1), but a studied plant temperature (t2). The
equivalent relative air humidity is expressed in this equation:

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 et1
rek   100
et 2

The faster the transpiration, the lower the relative air humidity and the higher
the deficit of vapour. When the relative air humidity (10–20%) is very low,
transpiration almost stops because of closing of stomata.

8.4.7.3.2 Temperature

A rising temperature brings about a rise in transpiration. For instance, vapour

pressure in the air at the temperature of 0 °C is 4.58 mm Hg, 9.2 mm Hg at 10 °C
and 31.3 mm Hg at 30 °C. If the relative air humidity is 50%, vapour pressure deficit
in the above temperatures is 4.6 mm, 9.8 mm and 29.5 mm Hg, respectively. This
fact reflects the progress of transpiration in the night influenced by the fall in
temperatures. Leaves, being photosynthetic organs, absorb solar energy and are a
location of various metabolic processes that heat them up. This results in an
increased difference between vapour pressure in the saturated environment (in
a leaf) and in its colder surroundings. In particular, plants with larger leaf surfaces
(and darker colour) can rise their temperatures faster, which speeds up transpiration.
At the temperature of 30 °C and the same level of radiation and relative air humidity,
transpiration shall be 9 times more intensive than at the temperature of 0 °C.

8.4.7.3.3 The light

Light radiation affects transpiration both directly by heating leaves and other
organs, and indirectly, as light is an impulse for stomata to open, which leads to a
higher stomatal transpiration.
Heating due to a direct light radiation is more important to plants that a fall in vapour
pressure in the atmosphere. Transpiration slows down linearly with a decreasing
radiation intensity and a falling temperature. On the contrary, in case of an increased
relative air humidity and decreased light intensity, transpiration rises.

8.4.7.3.4 The wind

The wind affects transpiration by removing vapour from the border layer of the
atmosphere above the leaf surface which may increase its water deficit. The value
of water potential in leaves falls due to the wind, which results in a disorder of a
formation of dry matter. A permanent air move brings about a formation of xeromorph
leaf structures. Plants get lignified and air holes get smaller.

8.4.7.3.5 The soil and other factors

This group of factors includes soil composition, soil humidity and content of
nutrients, air and gases in the soil. As stated above, not all water contained in the
soil is available to plants. The most suitable water capacity in the soil is 60% of the
full water capacity in the soil in terms of water available to a plant.
Transpiration in more fertile soils is lower. Even artificial fertilization can
decrease transpiration. After taking nutrients up, a plant changes its osmotic-colloid

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characteristics of the cell content. Absorbed nutrients play a part in the control over
the content of different forms of water in a plant. Nitrogen fertilization usually slows
down transpiration temporarily, but the subsequent growth of the foliage area speeds
transpiration up. Phosphoric acid fertilizers decrease transpiration as potassium
fertilizers do. Calcium affects various plant species differently, sometimes leading to
a rise, other times to a fall in transpiration.
The process of transpiration is greatly affected by the temperature of the soil
solution, the content of oxygen and other gases in the soil that are secondary
factors. They either slow down or speed up the uptake of water by roots, which
influence transpiration secondarily.
Exogenous factors of transpiration include emission, smoke gases and quakes.
Fly ash increase transpiration secondarily – as it falls on a leaf surface, it brings
about a corrosion of the cuticle or a grave dehydration of leaf tissues, thus increasing
transpiration. Quakes also bring about the increase in transpiration. This means it is
necessary to transport plants with a proper care.

8.4.7.4 Quantities of transpiration

The greatest consumption of water for transpiration occurs in late spring when
woody species have developed a foliage surface and the majority of leaves have
matured for photosynthesis. Moreover, this is a season of the quickest growth and
longest daytime. In central Europe, this season dates from May till late June,
depending on a species.
On a sun shining day, woody plants transpire 5–10 g of water per 1 m2 of foliage area
per an hour. Conifers transpire only about 1/10 of this volume by deciduous trees. A
beech forest (1 ha) transpires from 25.000 to 30.000 of water daily, and 3.600.000 kg
of water in a growing season.
The transpiration capability (i.e. rate between the amount of transpired water by a
plant and the potential evaporation – at the same place and time, or relative
transpiration) and maximal transpiration (i.e. average maximal speed of
transpiration measured at a natural site) are necessary quantities for the
estimation of water consumption by individuals of the same species at
different sites. The amount of water (e.g. in litres or tons) consumed for a plant or
stand transpiration in a growing season per a weight unit of dry matter (kg, t), is
called a transpiration coefficient, or the inverse value, i.e. efficiency of water
utilization, known as transpiration efficiency. Water turnover is a period of the
exchange of water in a plant (by means of transpiration and uptake of water). A plant
uses only about 0.01–0.08% of the total amount of transpired water for the creation
of a dry matter. The absolute majority of transpired water only serves as a medium
for nutrients and other substances transported in a plant body and as a cooling
system.

8.4.7.5 Anti-transpiration substances

A decrease in transpiration means a change in related physiological biochemical

processes. Anti-transpiration substances help decrease transpiration of 60–
80%, while leaf temperature rises of several degrees. Anti-transpiration
substances also control the uptake and transport of ions.
The main reason for the application of anti-transpiration substances is to decrease
transpiration at a certain period, thus protect plants from a lack of water. Yet, it is

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not clear if anti-transpiration preparations do not limit the entry of carbon
dioxide into a leaf, which would subsequently affect the photosynthesis.
Experience in the application of various anti-transpiration preparations shows that the
application is expensive and is not as efficient in numerous cases as was expected.
The anti-transpiration substances are recommended to be applied prior to
transporting during transplanting in a growing season, as they improve the success of
transplanting. Transpiration may be as well weakened by the increased content of
carbon dioxide in the air (greenhouses) or by the application of some growth
inhibition preparates (abscisic acid etc.).

8.4.8 Water balance in a plant

We distinguish between an active and a passive water balance. The active

(positive) water balance is a state when a woody plant plays an active part in water
saturation. The passive (negative) water balance means an increase in the water
saturation deficit and a disorder in water management.
The optimal water balance is a state when a plant does not suffer either from a
lack or a surplus of water, meaning that the activity of roots, rate of flow and
transpiration are maximal. As soon as the balance in the receipt, conduct and release
of water by a plant is disturbed, a greater water deficit brings about withering, related
with the accumulation of abscisic acid and a decrease in the content of gibberellins in
leaves. There are the following stages of withering: beginning, temporary,
permanent, irreversible.
The beginning stage of withering occurs to woody plants in hot summer days when
roots do not have enough capacity to store above ground parts with such a high
amount of water that is transpired due to high temperatures of the air.
Another problems may include the elimination of conductive pathways due to
entry of air from neighbouring tissues (formation of air bubbles and
embolisms), or blocking of conductive pathways by the growings from
surrounding parenchyma cells, thyllas. Especially round porose woody species
incline to thylas and embolizations. Conductive pathways of woody species may be
made dysfunctional also by parasitic fungi or damaged by phytofagous insects.
In the night, there is a decrease in air temperatures and a lack of available water in
soil, so a plant replaces the lost water, in the morning it is saturated with water and
appears fresh – turgescent. A greater water deficit in a plant results in a fall in
carbon dioxide uptake and limitation of photosynthesis due to closing of
stomata. In lack of available water in the soil or in hypoxia (lack of oxygen due to
compression of flooding) or salting, the beginning temporary withering usually passes
into permanent irreversible changes, which brings about a deep and permanent
damage that is reflected by accumulation of toxically substances, changes in
growth and gradual dying.

8.5 Mineral nutrition of woody plants and the significance of nutrients

Plant nutrition, i.e. substances gained from the neighbourhood and assimilated
substances in the body, is mineral in autotrophic species, organic in heterotrophic
and both mineral and organic in mixotrophic species. The nutrition is received by
body surfaces, which are really extensive in plant bodies compared to their volume.
In woody dry matter (with no water), there is approximately 45% of carbon, 42% of
oxygen, 6.5% of hydrogen and 1.5% of nitrogen. Carbon is taken in the form of

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gaseous CO2 from the atmosphere (besides a negligible amount in a liquid stage
taken up by roots from soil solution), oxygen originates from CO2 and hydroxyl ion
(OH–), hydrogen is from water molecules (H2O) and nitrogen along with the
remaining 5% of mineral substances from the soil.
Individual elements in a fresh weight of plants are represented in the following
rates: P, K,Ca, Si, S, Mg, Fe, Na, Cl, Al in 10–1 to 10–2%, known as
macroelements; Mn, B, Sr, Cu, Ti, Zn, Li, Ba, Br in 10–3 to 10–4 %, i.e.
microelements and traces As, Mo, Co, I, Ge, Pt etc in 10–5, ultramicroelements.
Ash elements are inorganic – mineral elements that remain from the dry matter of a
plant body after burning. Burning releases carbon and nitrogen from organic
bonds, both of them in the form of oxides. Most ashes are included in bark and
leaves (3–8% of dry matter), least of them is in wood (0.4–0.5%).
The above named mineral nutrients are also classified by the rate of the necessity
(physiological effect) and by the form they are absorbed – as cations, anions or
heavy metals. The ones with life significance are the nutrients without which a
plant is not able to enclose its life cycle, i.e. cannot give rise to germination seeds
and a new healthy generation. Moreover, these elements are not replaceable by any
others. These are biogenous elements with building or control functions –
catalysers, cofactors, moderators. By the general significance and content, they are
divided into macrobiogenous (anions: C, O, N, P, S, cations : H, K, Ca, Mg, heavy
metals : Mn, Fe, Cu, Zn) elements and elements in a smaller, or even trace
content, called oligo-biogenous elements (B, Cl, V, Mo, Bi, Ti, Ba, Li, Co etc.).
Qualitative requirements for the nutrition are more or less same in all green plants.
They differ in quantitative requirements that mainly reflect in different mutual rates.
While imbalanced nutrition, mainly by oligo-biogenous elements hardly exists in
natural conditions, human activities (by growing of single species and same age
stands, improper fertilizations and irrigations, chemical preparations in winter,
emissions, increase of CO2 and NOx in the atmosphere) violate the nutrition both by
biogenous and oligo-biogenous elements.
The source of nutrition is mainly in the solid phase of soil that releases elements
in the liquid phase to move to roots by diffusion or mass flow – rain water or
transpiration. Root exudates help make them more available (hydrogen cations, acid
carbon anions and organic acids). Then, individual nutrients enter roots via diffusion,
absorption exchange for excluded cations and anions and further on actively,
with the assistance of macromolecular protein transmitters. So a plant needs to
give out energy (ATP) gained in respiration of root cells in order to take in nutrients
into roots. This means that the uptake of nutrients needed to satisfy the following
needs: satisfactory temperature regime, sufficient water and oxygen in soil,
photosynthates in roots that become sources of energy in respiration. Healthy
nutrition of trees is also influenced by the volume of soil utilizable by roots and
capabilities of an individual to form, extend and maintain a sufficient sorption
area (surface of fine roots and root tips).
Another source of nutrients is their recoverability – e.g. in the fall of died plant
bodies (leaves, bark, roots) that are decomposed by soil organisms, releasing
mineral nutrition for a new usage. The recoverability of nutrients is also active by
secretion and excretion of plant elements, and to a little extent also by washing
away of easily mobile ions from leaves. Leaves that are lower positioned in a crown
of a tree can receive ions washed away from leaves positioned above them. This
makes a little receipt of nutrients by leaves. Leaves secure the carbon nutrition
(control element of quantity of plant production), though they are able to receipt even

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some ions and molecules through stomata or injured cuticle, e.g. ions of potassium,
sulphates, nitrites and nitrates, saccharose (i.e. nutrition else that in roots that is
successfully applied in intense agricultural fields and plantations of fruit trees as leaf
fertilization, or, on the other hand, entry of toxic elements into plants through the
barrier of covering tissues of a stem system).
Though, the main role in the woody plant nutrition is played by the root system,
not only in the uptake of nutrients, but also in the control of storage,
mobilization and separation into the above ground system.

8.5.1 Roles of individual nutrients

The roles of individual nutrients can be briefly described as follows: carbon, oxygen
and hydrogen (C, O, H) are the building stones of saccharides, organic acids and
lipids, while carbon is (almost) only accepted by leaves in the form of CO2 and
hydrogen comes from water taken in by roots.
Nitrogen (N) joins these three elements during the synthesis of amino-acids (some
of which contain sulphur). The uptake is active from the soil; in pH up to 6.8 it
enters root in the form of NO3– and its incorporation needs an active participation of
nitrate reductase and hydroxylamine reductase. The intake in the form of NH4+
dominates from pH 7, and it may be toxic in alkaline environment (it reduces intake of
cations, and even causes the release of K+ and Na+ from roots). The incorporation of
this form into aminoacids causes a release of H+ and may lead to a rise in acidity of
the cytoplasm. This danger is serious, particularly in urban coniferous stands as a
result of a high number of irresponsible dog keepers. Nitrogen may also be
absorbed in oxides that originate in the atmosphere during electric discharges
(including combustion engines) and that are dissolved in rain water. They could be
absorbed by leaves and roots. A higher nitrogen nutrition launches growth, limits
early maturation of wood and makes woody plants more sensitive to parasitic fungi
and unstable temperatures, especially low temperatures.
Phosphor (P) is bound in nucleoproteins and phospholipids. It is essential in
energy needs of metabolism (phosphorilation of saccharides, energy containers
of ATP, ADP – adenosine triphosphate, adenosine diphosphate, etc.). It is receipted
as H3PO4 or PO43– . The intake grows in the presence of calcium cations and
borates (synergy, see below). Phosphor transport and redistribution takes place in
phloem sections of vascular bundles.
Sulphur (S) is absorbed in the form of sulphates and stimulates the intake of
nitrates. Its way into the apoplast is passive, and it is transported into the symplast
by a specific permease built in the cytoplasm membrane. It is quite soon, already in
roots, that suplhur incroporates into specific proteins (cystein and glutation),
constituents of the antioxidation system.
Potassium (K) enters a plant as a potassium univalent cation as an exchange for a
hydrogen cation released into the rhizosphere by root cells. It can activate about 60
enzymes in the respiration process, utilization of energy, reduction of nitrates, protein
syntheses and reserve saccharides. It plays a significant part in the hydratation of
tissues and is released from cells in stress.
Sodium (Na) is taken by plants in less quantity than potassium, it accumulates in
vacuoles and acts as osmotic agent in halophytes – plants growing in saline
environment (mangrove swamps). Though, the absolute majority of woody plants

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control water movement in body by means of osmotically active saccharides or by
means of other simple organic substances (e.g. proline proteins etc.).
Calcium (Ca) participates in cell hydration by stimulating the uptake of K+,
stabilizing membranes and eliminating the permeability of these, and by eliminating
the release of ions, it can also activate certain enzymes, control elongation as well as
diameter growth and it takes part in the preservation of a good state of cell environs
through detoxication of organic acids. A considerably high amount of Ca is
contained by cell walls (Ca-pectates), while its concentration in cell cytoplasm is very
low.
Magnesium (Mg) is a constituent of middle lamellas in cell walls, of chlorophyll
molecules (as a central atom of porphyrin complex) and cooperates on the formation
of granas in chloroplasts. It activates Rubisco (one of the most significant enzymes
of photosynthesis) and DNA-polymerase enzyme. It is a part of some proteins, it
controls water regime and phosphate transport.
Iron (Fe) presents a transition to oligobiogenous elements, it occurs in the form of
organic (chelate) bonds, it is essential for the synthesis of chlorophyll and
reduction oxidation processes in metabolism.
Boron (B) is needed for the metabolism and transport of saccharides and
phenols, it plays a part in the elongation of pollen tubes and root cells, and,
above all, in the synthesis of nucleic acids. Moreover, the presence of boron (in
relation with the transport of phenolic substances) decides over the accumulation of
lignin and other defence structures.
Manganese (Mn) in enzymes acts in the process of oxidation (namely in
respiration), nitrates reductions, control of the rate of divalent and trivalent iron
and in the processes of photosynthesis.
Copper (Cu) stabilizes chlorphyll (through the influence of protein synthesis of
chloroplastic structures), it is a constituent of enzymes, it affects nitrogen
metabolism. In the form of oxides and phenolases, it takes part in tissues
lignification.
Molybdenum (Mb) is a constituent of nitrate reductase, and it supports the
assimilation (fixation) of nitrogen, required by soil microorganisms.
Zinc (Zn) has a range of functions in the oxidative metabolism and hydrolytic
reactions, it affects the assimilation of saccharides, biosynthesis of proteins
and some growth elements.

8.5.2 Ion relations

The ion antagonism is a phenomenon in which parallel factors generate opposite

effects. In case of mineral nutrition this means that one ion eliminates the entry of
another ion into a plant. E.g. an increased concentration of K+ and Cl– in the
neighbourhood leads to a limited uptake of Ca2+, Mg2+, Na+, but also of phosphorus
and sulphur. There are more antagonistic relations between ions, such as between
magnesium and calcium, phosphorus and calcium, sodium and calcium, between
forms of aluminium and iron and manganese and iron.
Moreover, the principle of synergism is applied in the uptake of ions. This means
that a two parallel factors have greater influence than if they were affecting
individually. For instance, the increased intake of nitrogen (NH4+, NO3–) or calcium
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(Ca2+) stimulates the intake of potassium (K+). Potassium ions have a positive effect
on the phosphorus intake.
On the other hand, the synergism may have a negative affecting, as well. Sulphur
dioxide and nitrogen oxides, if present along with ground ozone, cause more serious
damage to woody plants than if they acted individually.
Besides the ion relations, the nutrition intake also depends on the ion form in the
soil. E.g. K+ from potassium sulphate (K2SO4) is better accepted than K+ from
potassium chloride (KCl). Nutrition changes may be also caused by the
competitiveness of ions that is reflected in confusing a ion needed for nutrition with
another one of a similar structure. Such a relation may be observed with potassium
and rubidium ions. They both share the same active binding centres of
transmitting; so some centres are blocked by rubidium and cannot take a needed
amount of potassium. In another case, if significant respiration pathways are blocked
by an inhibitor, root cells may lack energy for an active distribution of needed
nutrients through barriers of cell membranes.
Another nutrition aspect is a plant capability of utilization of the absorbed
nutrition (e.g. to build it in a corresponding structure, enzyme etc.), or translocate it
into places with higher demands and reutilize one ion. On the other hand, a plant
needs to be capable of immobilizing nutrition absorbed in a surplus amount and
detoxication of some toxic substances (condensing of calcium ions and formation of
oxalate crystals or carbon calcid, chelating of heavy metals etc.).

8.5.3 Nutrition disorders

Nutrition disorders include not only insufficient nutrition by some elements

(differentiation), but also surplus nutrition (luxuriant) and imbalanced nutrition.
Trees react to nutrition disorders by changes in growth and its timing, but if the
disorders are more serious, it may lead to disease and visible damage to a plant, and
these changes (e.g. colour of leaves, changes in the development of certain
structures etc.) are evaluated as a deficiency disease.
For the adequate plant nutrition it is necessary that the intake of elements, their
transport (complicated in some elements due to immobility of these) and utilization
are not disrupted. Therefore the standard of mineral nutrition (e.g. availability and
usability of nutrients) needs to be studied as a whole from various angles. This
means to analyse it from the perspective of soil conditions and its physical chemical
characteristics, and from the perspective of the plant which is complex since the
occurrence of substances, amount of substances, form and mutual ratio of
substances varies by structure, species, ontogenetic stage, season. All these factors
show that even a fertilization decision which seems easy at first sight may be a
contra-productive action.

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