(A) Medullary Ray in in the Wood

木材中的髓放線

(B) 1. Ring porous 環孔材

[Link] porous 散孔材
Concerning the wood of a tree, there are
differences between the wood of conif-
erous and deciduous trees. The wood of
deciduous trees has further differences
Concerning the wood of a tree, there are
differences between the wood of conif-
erous and deciduous trees. The wood of
deciduous trees has further differences
between ring-porous wood (i.e. Quercus,
Fraxinus) and diffuse-porous wood (ie.
Tilia, Fagus).
Heartwood is formed within the centre
of the tree. The heartwood is surrounded
by a ring of living sapwood.
Not all of the conducting elements in the
xylem transport water. Only those of the
sapwood (active and living wood) func-
tion to conduct water, while the other
elements of the xylem which are situated
farther inside the tree are no longer
functional. 'They constitute the heart-
wood, non-conducting tissue that is
sometimes darker in colour than the sap-
wood. The phloem is responsible for the
movement of sugars, produced in the
leaves, to other parts of the plant.
In addition to the phicem and xylem, the
vascular system of a tree contains ray
cells. Rays grow out radially from the
centre of the tree to cross through the
phloem and xylem and function to trans-
port sugars and other compounds across
the trunk. They assist in restricting decay
in wood tissue and store reserves in the
form of starch.
The outer covering of a tree's branches
and stems is the bark. It is a protective

tissue that moderates the temperature

inside the stem, offers defence against
injury, and reduces water loss. Bark is
composed of non-functional phloem,
corky tissues and dead cells. The cell
walls are impregnated with wax and oil
that minimize water loss. Lenticels, small
openings in the bark, allow for gas
exchange between the living tissues of
the tree and the atmosphere. Many
types of bark develop in trees. For
example, Fagus has very smooth bark.
with little corky material and Quercus
suber produces thick layers of cork.
Branches and Twigs
Twigs are small stems that provide the
support structure for leaves, flowers and
fruit. Branches support twigs, and the
trunk supports the entire crown.
Branches and twigs emerge from two
types of buds:
- the terminal or apical bud at the end
of a shoot and
- the lateral or axillary bud occurs along
the twig.
The terminal bud is the most powerful
bud on the branch or twig and located
at the end of a shoot. It controls the
development of the secondary buds
by means of hormones. Usually the sec-
ondary buds do not develop and are dor-
mant.
Normally, the terminal bud is the most
active on each branch or twig and con-
trols the development of the lateral buds
on the same shoot which are often dor-
mant: Their growth is inhibited by the
apical dominance of the terminal bud.
Apically dominant shoots are monopo-
dial or sympodial. Shoots that are not
apically dominant are pseudodichoto-
mial. The destruction of the terminal bud
by accident or pruning can release dor-
mant buds near the cut, leading to new

Branch Formation 分枝類型

MONOPODIAL SYMPODIAL PSEUDO DICHOTOMY)
單軸分枝 合軸分枝 假二叉

Annual Growth 年度生長變化

shoot development. Some shoots
develop from adventitious buds which
are produced along stems or roots. They
are usually produced in response to the
loss of normal buds and the growth reg.
ulators produced by them.
Leaves and buds are from a slightly
enlarged portion of the twig called a
node. The internode is the area between
the nodes. Leaf scars and terminal bud
scale scars are visible on the twig. These
are useful in measuring annual twig
elongation or growth.
Each branch of the tree is similar in struc-
ture and function to the entire tree
crown. Yet branches are not simply out-

growths of the trunk. Instead, branches

have a unique attachment form, which is
critical to the application of arboricultural
practices such as pruning. Branches are
strongly attached to the wood and bark
beneath the branch but weakly attached
to the wood and bark above the branch.
The annual production of layers of
tissue at the junction of the branch to
the stem is apparent, forming, most of
the time, a shoulder or bulge around the
branch base called the branch
collar.
In the crotch, the branch and trunk
expand against each other. As a result,
bark is pushed up forming the branch
bark ridge. If bark in the crotch is sur-
rounded by wood, it is called included
bark. Included bark further weakens the
crotch since the normal branch to trunk
attachment is not formed.
分枝連接

A - 莖 / 活分枝
B - 莖 / 死分枝

1. 莖材
2. 活木
3. 死木
4. 保護區
5. 新莖材

growths of the trunk. Instead, branches

have a unique attachment form, which is
critical to the application of arboricultural
practices such as pruning. Branches are
strongly attached to the wood and bark
beneath the branch but weakly attached
to the wood and bark above the branch.
The annual production of layers of
tissue at the junction of the branch to
the stem is apparent, forming, most of
the time, a shoulder or bulge around the
branch base called the branch
collar.
In the crotch, the branch and trunk
expand against each other. As a result,
bark is pushed up forming the branch
bark ridge. If bark in the crotch is sur-
rounded by wood, it is called included
bark. Included bark further weakens the
crotch since the normal branch to trunk
attachment is not formed.
葉片構造
1.氣孔
2.角質層
3.表皮
4.柵狀薄壁細胞
5.海綿薄壁細胞
Leaves
Leaves are the food producers of the
tree. They contain chloroplasts that
are filled with a green pigment called
chlorophyll, where photosynthesis
occurs.
A second role of leaves is transpiration.
Transpiration is the loss of water through
the foliage in the form of water vapour.
Leaves provide a large surface area for
the absorption of sunlight and carbon
dioxide needed for photosynthesis. The
outer surface of a leaf is covered by a
waxy layer called the cuticle. The cuticle
functions to minimize dessication (drying
out) of the leaf. Stomata, small, variable
openings in the leaf surface, control the
loss of water vapour and the exchange
of gases.
Leaves have a network of conducting
tissues that comprise the veins, or
vascular bundles. These veins are
composed of both phloem and xylem
tissue. They transport water and essential
elements, and carry food produced in
the leaf cells to other parts of the tree.
Trees that shed their leaves every year.
are called deciduous. Trees that retain
their leaves for more than one year are
called coniferous or evergreen. Leaf drop
is caused by cell changes and growth
regulators that combine to form an
abscission zone at the base of the leaf
stalk or petiole.
The abscission zone has two functions:
- to enable leat drop in the autumn
- to protect the region of the stem from
which the leaf has fallen against
dessication, insect or disease damage.
Autumn foliage colour in deciduous trees
results from the breakdown of chloro-
phyll to reveal other pigments contained
in the leaf. Shorter days combined with
cold nights enhance the accumulation of
sugars and trigger a decrease in
chlorophyll production. This allows other
pigments including anthocyanins (reds
and purples) and carotenoids (yellows,
oranges and reds) to be unmasked.

Roots
The roots of trees serve four primary
functions:
- anchorage
- storage of energy
- absorption
- conduction
Absorbing roots are the small, fibrous,
primary tissues that grow at the ends of
the main, woody roots. The absorbing
roots have epidermal cells that are
modified into root hairs, which aid in the
uptake of water and minerals.
The root hairs only have a short life
(3-4 weeks in spring) and duplicate
considerably the possibilities of absorp-
tion when vegetation starts in spring.
As with shoot tips, root tips contain
a meristematic zone, where the cells
divide and grow in length.
Roots grow where moisture and oxygen
are available. Most absorbing roots are
found in the upper 30 cm of soll.
Horizontal, lateral roots are also usually
A. 根系

1. 主根系 2.心形根系 3.表面根系

B.根梢

1.木質根 2. 根毛 3. 根尖 4.根冠

near the soil surface. Sinker roots grow

vertically downward off the lateral roots,
providing anchorage and increasing the
depth of soil exploited by the root
system.
Many roots live in a symbiotic relation-
ship with certain fungi. The result of the
association is termed mycorrhizae (fun-
gus roots). In symbiosis, both organisms,
the tree and the fungus in this case,
benefit from the living arrangement. The
fungi derive nourishment from the roots
of the tree. In turn, the fungi aid the
roots in the absorption of water and
essential mineral elements.
光合作用基本要素
水 (H₂O)
糖（葡萄糖, C₆H₁₂O₆）
氧氣 (O₂)
二氧化碳 (CO₂)
太陽能（光能）
蒸散作用

2.1.3 Tree Physiology

Photosynthesis
Photosynthesis is the process by which
green plants use light energy to build
sugar molecules. This reaction takes
place within cells that contain chloro-
plasts. Chloroplasts contain molecules
of the light-absorbing pigment, chloro-
phyll. Chlorophyll gives plants their
green colour.
The raw materials necessary for photo-
synthesis are, apart from light energy,
carbon dioxide and water. The tree
absorbs carbon dioxide from the at-
mosphere through the stomata in the
leaves and water is drawn from the
soil. Light energy is absorbed and
trapped in the chloroplasts. The light
energy is converted to chemical energy
and stored in the form of sugars and
starches. Oxygen, the by-product of
photosynthesis, is released through the
stomata. The sugar products of photo-
synthesis are the building blocks for
many other compounds required by the
plant. Proteins, starch, fat, vitamins,
amino acids and other important
compounds are produced from sugar
when combined with other essential
elements such as nitrogen, potassium,
sulphur and iron.
Respiration
Respiration is the oxidation of organic
molecules, a process by which the
chemical energy generated by photo-
synthesis, and stored as starch or sugar,
is used by the tree. Respiration that
occurs during the hours of darkness us-
es the energy produced during the day
by photosynthesis.
When respiration takes place in the
absence of photosynthesis, the tree
must rely on stored energy reserves.
If this occurs over a long period of
time, the tree will eventually run out of
energy and die. A practical example is a
tree that is repeatedly defoliated. With-
out foliage, photosynthesis stops, and
the tree cannot produce and store
starches. Yet as long as the tree is
living, respiration takes place, and
energy is consumed.
Transpiration
Transpiration is the loss of water in the
form of water vapour principally from
the underside of the leaf surfaces. The
cuticle helps prevent uncontrolled water
loss from epidermal cells on the leaf sur-
face. Gas exchange, where water vapour
and oxygen are released and carbon
dioxide is absorbed, takes place through
stomata, also on the underside of the
leaf. Each stomata is lined by two guard
cells, which regulate the opening and
closing of the stomata. Stomatal opening
is influenced by environmental
conditions such as light, temperature
and humidity. Stomata usually open in
the light and close in the dark.
The rate of transpiration is affected by
temperature, humidity and available
water. Transpirational water loss is also
affected by anatomical features such as
cuticle thickness, the presence of hairs
on the leaf surface, and the number and
ocation of stomata.

To a lesser degree transpiration also

takes places through the living wood
and bark. Through the bark the living
tissue in the wood absorbs oxygen and
releases carbon dioxide. The tree can
also dessicate, by means of water
evaporation, even after leaf fall when
the tree is not in contact with the soil -
for example on a planting site during
pring and autumn or whilst being trans-
ported.
蒸散作用（Transpiration）在較小程度上也會通過活
木（Living Wood）和樹皮（Bark）進行。通過樹皮，
木材中的活組織吸收氧氣並釋放二氧化碳。即使樹木
在落葉後不與土壤接觸（例如在春季和秋季的種植地
點或運輸過程中）
，樹木仍然可以通過水分蒸發而乾
燥（Dessicate）。
Water and Nutrient Uptake
Water and essential elements are
principally absorbed from the soil by
the roots. To a lesser degree this also
takes place through the leaves.
Some of this water is used by the tree
for growth and metabolism, but most
is lost through transpiration. This water
loss creates the , transpirational pull"
that, among other processes,
moves water through the xylem.
Water enters young roots by a process
called osmosis. Osmosis is the
movement of water through a mem-
brane from a region of high potential
(water concentration) to a region of
low water potential. Water normally
will move into roots where the water
potential is lower than in the surround-
ing soil. If the water potential is lower

in the soil than the root cells, water will

actually move out of the roots into the
soil (exosmosis). An example is when
salt concentrations are high in the soil
such as from deicing or excessive
fertilizer application.
The sugar products of photosynthesis
are actively pumped through the
phloem, a process which requires a lot
of energy.
The transportation of water and sugar
basically takes place in all parts of the
tree.
Growth and Development
The growth and development of a tree
is the result of the interaction between
its genetic potential and the surround-
ing environmental conditions. The
urban environment often limits the
expression of that potential, especially
the range of size.
A tree is particularly complex in
structure. From the architectural point
of view, the tree is a genetically deter-
mined system of branches, stems and
roots, each element deriving from one
another by ramification. Each branch of
the plant possesses a particular
function and structure which derives
from its order in the ramified structure.
A young tree possesses a homogenous
functional architecture that derives from
its biological basic structure. So a young
plant is formed as a single architectural
unit. Gradually, the organism multiplies
the number of its units. They become
smaller and smaller and thus become
more difficult to analize. The analysis of
the main branches of an older tree
shows that each branch is a miniature
tree or a collection of small ramified
trees called reiterates.
The development of a tree can be
described in three main phases:
- the juvenile phase,
- the stage of maturity,
- the regression phase.

發展階段
幼年期
中年期
成熟期
Each stage has its own physiological
characteristic and growth.
The tree permanently increases in
diameter (stem and crown), initially
rapidly, especially the growth of its
height, but slowing with age. The
mature, adult tree is totally functional.
The stage of regression is between the
decreasing vitality and the death of the
The coordination of growth processes
in trees is controlled in part by plant
growth substances or hormones.
Hormones control cell division, cell
elongation, fruit ripening, leaf drop
and root development.
Auxin is one of the most important
plant growth substances. It is known to
be especially important in root develop-
ment. Synthetic forms of auxin are sold
commercially to enhance the rooting of
cuttings.
tree.

CODIT 模型（CODIT Model）

木材中的屏障區（Barrier Zone in Wood）
1 Wall 1
○
2. Wall 2
○
3 Wall 3
○
4 Wall 4
○
Compartmentalization
Compartmentalization is one of the
most important reactions of a tree to
damage or infection. This mechanism
enables trees to adapt their develop-
ment to the new situation by limiting
the spread of discolouration and decay.
After a tree has been wounded,
reactions are triggered which cause the
tree to form boundaries around the
wounded area. The boundaries are
called reaction zones.
Dr. Alex Shigo has proposed a model of
this compartmentalization process
called CODIT (Compartmentalization of
Decay in Trees). In this model, the tree
forms four barrier walls:
- Wall 1 resists vertical spread by
plugging xylem vessels
- Wall 2 resists inward spread by more
compact summerwood cells and by
depositing chemicals in these cells
- Wall 3 inhibits lateral spread by
activating ray cells to resist decay
- Wall 4 is the next layer of wood to
form, this time by the cambium,
after injury. This protects against the
outward spread of decay and is
called the barrier zone - the
strongest of the four walls.
The effectiveness of the different walls
depends on the vitality of the tree. It is
fairly common for walls 1, 2 and 3 to
fail, allowing decay to spread inside the
tree, forming a hollow cavity. Wall 4
rarely fails, except where canker-
causing fungi restrict its development
or kill the cambium.
The ability to compartmentalize varies
between the different tree species.
Examples of good or strong comparte-
mentalize species are:
Acer pseudoplatanus, Acer campestre,
Carpinus, Crataegus, Fagus, Gleditsia,
Platanus, Pinus, Quercus robur, Taxus,
Tilia [Link], Ulmus.
Examples of tree species that are weak
to compartementalize are: Abies, Acer
sacharinum, Aesculus, Betula, Fraxinus,
Malus, Picea, Populus, Prunus, Salix.

2.2
Tree/Soil Relations
2.2.1
Introduction
The relationship between tree root
systems and the characteristic of the
soils in which they grow has a greater
influence on tree health than any other
single factor. By understanding soil tex-
ture, structure, pH, and water-holding
capacity, the Tree Worker will be better
equipped to manage trees in an urban
environment.

Soil Horizon 地底水平線

2.2.2
Properties of Soil
[Link]
Physical Properties of Soil
Naturally occurring soils are the result
of thousands of years of biological,
chemical and physical weathering and
erosion of the parent material, or
underlying bedrock. Soils are usually
dominated by the geology of the
parent material.
The soil profile normally consists of
four major layers or horizons. The top
of the profile is a thin layer of decom-
posing organic material called the
organic layer. The next layer down is
the A horizon. The A horizon contains
most of the absorbing roots of trees.
This section is rich in organic matter.
The B horizon, intermediate in depth, is
composed of fine textured materials
from the A horizon and soil particles
from the lower parent material. The
A and B horizons are often mixed and
form the ,topsoil" as it is called in
connection with urban tree sites.
The C horizon is the lowest layer and is
composed of shattered and weathered
rocky parent material. The topmost
layers are very active biologically and
there the litter is decomposed by the
activity of micro-organisms. The litter
is composed of leaves, twigs, bark etc.
The tree roots grow where soil condi-