Tree Architecture and Growth

Conceptually, a tree is a tower supporting many small

solar collectors.
The objective is to get lots of solar collectors (leaves)
in the air with the least cost in photosynthate / carbon.
The design must be strong enough to last decades or
even centuries. Over-designing wastes materials
needed in other parts of the structure.
Unlike towers, trees are dynamic and grow, but growth
comes only from materials trees themselves produce.

Forest Stand Dynamics

Winter 2003

Primary and Secondary Growth

Stand development patterns emerge because trees have

innate growth habits and respond with characteristic growth
patterns to physical surroundings (site, weather,
disturbances) and to other plants competing for the same
growing space.

Primary growth - growth from a bud, root tip, or other apical

meristem

The shape of individual trees and whole stands is a key to

past history.

Meristems are localized areas that act as cell factories and cause
elongation, thickening, and branching - producing new cells by division;
these cells enlarge and differentiate into the many kinds of tissues that
make up a tree.

Trees add new growth to old structures that developed

under different conditions.
Tree shape is primarily determined by primary growth,
allowing new growth to respond to recent or current
environmental conditions.

Secondary growth - growth from cambial or lateral meristems

Apical meristems are found at the tips of each growing shoot and root
and are responsible for elongation and producing the cells that form new
meristems.
Lateral meristems are located in the cambium between the wood and
the bark; these meristems produce cells that thicken roots, stems, and
branches and cells that increase the size of the lateral meristem .

Primary and Secondary Growth

Primary and Secondary Growth

Roots and shoots grow additively - new cells are added that make them
longer or thicker; total size is the sum of these additions.

A sequence of cells, tissues, or organs at increasing distance

from the meristems that formed them represents a
developmental sequence in time.

Growth in TOTAL number of roots and shoots is multiplicative

because the rate is determined by the number of parent shoots
multiplied by the branching rate per shoot.
Branches are not formed randomly; branching pattern - phyllotaxy arises from the apical meristems and can be alternate (oak, hickory,
cottonwood), opposite (maple, ash, dogwood), or whorled (eastern
white pine, Douglas-fir, true firs).
Tree form arises from annual repeating patterns of branching,
elongation, thickening, and death.
Differences in form arise from different process RATES, not diff erent
processes.

A record of cambial activity is stored in the wood and a

record of annual elongations is found in the shoot system.
It is possible to work back in time to determine the
development of the tree over time (stem analysis,
dendrochronology ).
Ice storms, insects, diseases, management, etc. affect form;
trees react by trying to restore original growth patterns, but
evidence of past events often remains.

Annual rings are a key to past

climate and disturbances.
This tree established in 1685 and
was only an inch in diameter
when it survived its first fire.
Fire scars from this and nearby
trees show fires occurred at
intervals from 7 -14 years until
about 1906.
In 1994, after nearly 100 years of
fire exclusion, this and all nearby
trees were killed in the Tyee Fire
on the Wenatchee NF.

Design Problems of Trees and Some Solutions

Pistol Butt is
caused by soil
creep on steep
slopes or by snow
damage when tree
was a seedling or
sapling.

Gravity
Because of gravity (aided and abetted by wind, rain, and
snow) aerial parts of trees tend to bend down.
One solution is to orient the main trunk with respect to
gravity. This is termed geotropism. Most conifers are
strongly geotropic. The shoot goes up, the root goes down.

Design Problems of Trees and Some Solutions

Strength
When shoots go up very far, they face the problem of
developing sufficient strength to avoid breakage from
wind or their own weight

Design Problems of Trees and Some Solutions

Anchoring
Solutions - thicken roots near the base of the stem to make them
stronger; develop a deep taproot

Solution 1 - develop wood - strong walled cells

composed initially of cellulose, then lignin
Solution 2 - thicken stems and branches to make them
stronger

Design Problems of Trees and Some Solutions

Growth

Design Problems of Trees and Some Solutions

Reduced Photosynthetic Efficiency

Growth creates need for more photosynthate - which requires

more leaves

Adding additional leaves creates the tendency for some leaves to

shade others, reducing photosynthetic efficiency

Solution - branch to produce more leaf-bearing axes, giving an

exponential increase in leaf number

Solution - develop a branching pattern and leaf arrangement that

minimizes overlapping
Monolayer on outside of tree crown (good in low-light or at
low latitudes where sun is overhead)
Multi-layer distribution throughout crown (good at higher
light intensities or at high latitudes where sunlight comes
from the side)

Design Problems of Trees and Some Solutions

Shaded Branches

Design Problems of Trees and Some Solutions

Water Loss During Photosynthesis

As the tree grows, branches on the lower and inner parts of the
tree become shaded and produce less photosynthate.

Increasing the number of leaves increases water loss through

stomata when CO 2 is taken up during photosynthesis

Solution - Kill off branches that are not self-sustaining and

concentrate leaves at the edges of the crown unless the crown is
sparse and the tree is growing in full sunlight

Solution - Proportionally increase the number of fine roots to

take up water. Maintain feedback mechanisms between roots
and leaves so they remain in balance (basis of pipestem carbon
balance models)

Design Problems of Trees and Some Solutions

As the tree gets larger, more

water and nutrients need to move
up and there is more
photosynthate to transport down.
Solution - Continually create
more transport cells (the function
of the cambium which creates the
cells that differentiate into xylem
(to move water and nutrients up
the tree) and phloem (to move
photosynthate down))

Design Problems of Trees and Some Solutions

Age and Injury As the tree gets older, the probability of

serious injury increases.
Solution - Produce replacement shoots or
roots that grow in the same direction as the
broken one and replaces its function
a) express tiny shoots/roots that have been
stored for emergencies (short lag time)
b) grow new shoots/roots (number and
orientation not predetermined so can adapt
to specific injury)
c) change orientation of existing branches
(lag time decreased as new growth not
required)

Design Problems of Trees and Some

Solutions

And Theyre Still Working On

This One...

Shoot Development

Growth of the terminal shoot controls the trees

architecture by controlling
branch growth

400 or 120 m appears to be the maximum

tree height possible
internal water stress increases with tree
height

crown shape
stem growth
Shoot growth entails the development of primordia
from the apical meristem plus active shoot expansion.

increased height requires increased

thickening at the base, adding weight
and causing structural problems

Shoot Development

Primordia are small precursors of stems or leaves

formed within the apical dome before and during active
shoot elongation.
The top of the apical meristem is a dome.
Bulges form on the dome flanks and expand by cell
division to form leaf primordia.
As primordia develop, the dome expands upward,
adding younger primordia within and above older
ones (think of an artichoke.).
Older primordia expand during shoot elongation to
form leaves or needles.

Shoot Development

Shoot Development

Most woody plants have woody bud scales that protect

primordia during adverse seasons.

Elongation is triggered by environmental conditions

such as accumulated temperature (degree days, blossom
degree days), day length, moisture, or combinations of
these.

A few species (witch hazel) have naked buds.

Budbreak signals the period of active shoot elongation.
During shoot elongation the primordia expand rapidly,
develop stems, leaves, and lateral primary meristems
(generally shoot or flower buds).

Cessation of elongation can be triggered by drought or

photoperiod.
Many tropical species have periods of active elongation
and dormancy, generally triggered by moisture
availability.

Root Development

Root growth can occur throughout the year under

favorable conditions.
Root growth usually peaks in early spring prior to shoot
elongation and in late autumn after shoot elongation and
cambial activity cease (this is why early spring and
autumn are good times to plant landscape trees).
Different tree species are predisposed to different root
growth patterns, but soil conditions exert a strong
modifying force on root growth.

Preform or Fixed Growth

Tree Growth Patterns

There are about 23 - 26 generalized growth
patterns for woody plants.
Five growth patterns are common to
temperate forest tree species in the northern
hemisphere:
Preform or fixed growth
Neoform or sustained growth
Recurrent growth
Terminal florescence
Aborted tip (sympodial/zigzag)
growth

Preform or Fixed Growth

A pattern of bud development, dormancy, and

activation found in many trees that survive periods of
environmental extremes by going dormant.

At budbreak, the preformed primordia elongate, with

expansion limited by the number of primordial cells
formed the previous year.

New bud scales are formed from leaf primordia at tip

of new shoot before or during active elongation.

Examples of trees with the preformgrowth pattern

include oaks, true firs, Douglas-fir, hickories,
spruces, ashes, and some pines.

Apical meristem develops new leaf primordia PRIOR

to next years active growing season.
Primordia reflect conditions of year they are formed,
NOT the year they are expressed.

Trees with preformgrowth generally form distinctive

annual rings (ring porous) because of differences
between early and late wood.

Lammas Growth
A variant of pre- form growth where a second flush of
growth occurs in mid to late summer from buds that
would ordinarily flush the following spring.
This generally occurs when growing conditions have
been unusually favorable.
Lateral buds as well as terminal buds can develop as
lammas growth.
When a lateral bud flushes it can take over the
terminal position, leading to a crooked stem.

Neoform (Sustained )Growth

Not all primordia develop prior to active shoot
elongation.
A bud containing a small number of primordia is set
near the end of the growing season.
The following year, these primordia develop during
shoot elongation, with new primordia continuing to
develop and shoots elongating as long as conditions
are favorable for growth.

Neoform (Sustained) Growth

Shoot expansion is generally not as rapid as for
species with preformed growth but often continues
for a longer time.
Species with neoform growth are generally more sitesensitive than species with preformed growth.
Species having neoform growth include sweetgum,
hemlock, red alder, yellow poplar, and red maple.

Amount of growth mostly depends on environmental

conditions during the CURRENT year.

Diffuse Porous v Ring Porous Wood

Neoform (Sustained) Growth
Diffuse porous red maple

Trees with neoformgrowth produce less definitive

growth rings (diffuse porous) because there is little
difference between wood formed early in the season
and wood formed late in the growing season.

Recurrent Growth

Ring porous sassafras

Recurrent Growth

Typical of southern pines; superficially resembles preformed

growth.

Vigorous trees produce more growth flushes than

weaker trees.

An inner bud is formed at the top of an enclosed shoot prior

to the opening of the outer bud.

There are no annual whorls of branches; whorls occur

at the ends of the telescoping shoots and not at the
end of an entire years growth.

A preformed shoot with another inner bud is telescoped

inside the topmost inner bud.
Several preformed shoots and inner bud combinations can
be telescoped together.

Trees with this growth pattern often have false rings

in the annual xylem, making aging difficult.

At budbreak, each bud and shoot breaks and elongates in

sequence, but often before the previous shoot has fully
elongated.

Terminal Florescence

Aborted Tip Growth

The terminal shoot develops into a flower.

Also referred to as sympodial or zigzag growth.

The flower exerts control over lower branches,

causing them to grow more laterally, resulting in a
series of forking branches.

Trees with this pattern abort their terminal shoots,

leaving the lateral branches to resume growth.

Plants with this growth form

include horsechestnut, sumac,
devils club and many tropical
trees.

Aborted Tip Growth

Trees with this growth pattern include birches, elm,
redbud, and sugarberry.

This pattern is distinctive, but can derive from either

preformor neoformgrowth patterns.
Each leaf acts as a continuation of the stem, and the distal
part of the stem then grows at an angle.
Trees often die back during unfavorable summers, then
resume growth the following year.

More on Growth
Very young trees of many species exhibit neoform
growth, even those that as adults have preform growth
patterns.
Foxtail - develops when the terminal shoot
grows without producing any lateral
branches; this often occurs with temperate
climate pines grown in the tropics,
perhaps as a result of there not being
enough stimulus for the terminal shoot to
cease growth or produce lateral buds and
branches.

Apical Dominance, Apical Control, and

Epinasty: the Real Story

Apical dominance is the tendency for CURRENT year

lateral buds to remain dormant as the shoot expands.
Auxins (growth hormones) in the terminal bud flow
downward, inhibiting growth of proximal (current
year) lateral buds.
Auxin concentrations weaken as they flow downward,
so PREVIOUS years dormant lateral buds flush and
lateral branches grow vigorously.

Apical Dominance, Apical Control, and

Epinasty: the Real Story

Species with strong apical DOMINANCE (the ability

to keep current year lateral buds dormant) have weak
apical CONTROL (they are unable to control the
flushing and growth of prior year lateral buds)
The result is a tree with long, heavy, lateral
branches (oaks)

Apical Dominance, Apical Control, and

Epinasty: the Real Story
When there is weak apical control,
lateral branches can grow upward
and act as leaders, leading to a
decurrent or rounded growth
form.

Older trees often revert to a

decurrent growth form.

A tree with weak apical DOMINANCE is NOT able to

completely suppress the growth of current year lateral buds, so
they flush.
Auxins from these buds also move downward and and
RESTRICT growth on previous years flushed buds.
ALL the branches remain rather short since auxin from the
terminal bud keeps current year lateral branches short and
auxin from each years lateral branches keep prior years
lateral branches short.
The result is a columnar tree with short lateral branches
(Fir)

Apical Dominance, Apical Control, and

Epinasty: the Real Story

Apical control refers to the physiological condition

governing the excurrent (single stem, pyramidal) or
decurrent (branchy) pattern of growth.
A tree with strong apical control maintains a single
stem, while one with weak apical control has many
branches and a rounded crown.
The term apical control will be encountered
MUCH more frequently than the term apical
dominance.

Apical Dominance, Apical Control, and

Epinasty: the Real Story
Apical dominance determines whether or not a bud will
expand; apical control determines how much the bud
will grow once it does expand.
Strong apical dominance = weak apical control =
branchy tree
Weak apical dominance = strong apical control =
columnar tree
PREFORMED SPECIES DO NOT NECESSARILY
HAVE STRONG APICAL DOMINANCE.

Apical Dominance, Apical Control, and

Epinasty: the Real Story
The textbook uses the term epinastic control when
referring to the concept of apical control.
There is no such thing as epinastic control!
Epinasty refers to greater growth on the upper side of a
branch, causing it to bend downwards.
Trees growing in shade lose APICAL control: their
lateral branches grow longer relative to their terminal
leader.