publication 422-025

Physiology of Pruning Fruit Trees

Richard P. Marini, Extension Specialist, Horticulture; Virginia Tech

Introduction
Woody plants are pruned to maintain a desired size
and shape and to promote a certain type of growth.
Ornamental plants are pruned to improve the aesthetic quality of the plant, but fruit trees are pruned
to improve fruit quality by encouraging an appropriate balance between vegetative (wood) and
reproductive (fruiting) growth. Annual pruning of
fruit trees always reduces yield, but enhances fruit
quality. Pruning increases fruit size because excess
flower buds are removed and pruning encourages
the growth of new shoots with high-quality flower
buds. Pruning improves light penetration into the
canopy, and light is required for flower-bud development, fruit set and growth, and red color development. Pruning also makes the canopy more open
and improves pest control by allowing better spray
penetration into the tree; air movement throughout
the canopy is increased, which improves drying
conditions and reduces severity of many diseases.
This publication describes why plants respond to
pruning and other forms of plant manipulation used
to train trees. This information applies to all plants,
but application to fruit trees is emphasized.
Pruning fruit trees is somewhat of an art based on
an understanding of plant physiology and development. In other words, if we understand how plants
grow and how they will respond to different types
of plant manipulations, we can alter vegetative
growth and fruiting to obtain trees and fruit with
desirable characteristics.
A basic understanding of certain aspects of plant
physiology is a prerequisite to understanding pruning. Unlike animals, plants continue to increase
in size throughout their lives. There are only two

ways plants can grow.

Primary growth is the increase in length of shoots
and roots, and is responsible for increases in canopy height and width.
Secondary growth is the increase in thickness of
stems and roots.
Both types of growth require cell division followed
by cell enlargement and differentiation.

Plant Growth
Meristems are regions of cell division and there
are two types of plant meristems. An apical meristem is located at the tip of every shoot and root
(Figure 1). As cells divide in these apical meristems, the shoots and roots elongate as cells are
piled one on another. Behind the region of cell
division is a region of cell differentiation, where
cells enlarge and differentiate into various tissues.
In the axil of each leaf is a small apical meristem
called an axillary meristem that forms an axillary
bud, which usually remains dormant until well
after the subtending leaf is fully developed. An
axillary bud may remain dormant or develop into
a lateral branch or a flower.
There are two distinct layers of meristematic tissue
within the stem or root responsible for secondary
growth, the vascular cambium and the cork cambium (Figure 2). The vascular cambium is a cylinder of specialized cells, usually five to ten cells
thick, running the length of the plant, including the
roots, and is responsible for the radial growth of
plant parts. Phloem cells are produced to the outside of the cambium and xylem cells are produced
to the inside of the cambium.

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vascular
cambium

Leaf
primordia

Apical
meristem

phellogen

bumps
phloem

xylem

periderm
(bark)

Figure 2. Longitudinal cross-section of a tree trunk shows

the vascular and cork cambiums.

axillary
buds

Figure 1. Longitudinal section of shoot tip shows an apical

meristem, successively older leaf primordia, and axillary
bud primordia.

a vegetative bud is sliced longitudinally during

the winter and viewed under magnification, the
apical meristem at the tip, leaf primordial (developing leaves), axillary meristems, developing axillary buds, and procambial tissue (tissue that will
develop into the cambium) are all visible.
Buds on fruit trees usually have about seven
leaves and initial shoot elongation in the spring
results from cell expansion. During late June and
July some of the shoot apices will flatten out and
develop into flower buds. Flower buds are actually modified shoots and the various flower tissues
(petals, stigmas, anthers, etc.) are actually modified leaves. Although the process of switching
from vegetative to reproductive buds is not fully
understood, hormones that can be influenced by
environmental factors, stresses, and plant nutrition
control the process.
There are several things we can do to influence
whether or not a bud becomes a flower bud or
remains vegetative. In general, factors that favor
rapid growth, such as high nitrogen levels in the
shoot tissues, inhibit the development of flower
buds. Applying growth-promoting plant growth
regulators such as gibberellins usually inhibits
flower-bud induction, whereas ethylene may promote flower-bud development. Mild stresses such
as shoot bending and water stress may also promote flower-bud development.
Producing annual crops of high-quality fruit
requires a balance between reproductive and vegetative growth. Fruit producers use various techniques, including pruning, branch bending, and

Downward transport of sugars, nutrients, and hormones from the top of the tree to the roots occurs
in the phloem tissue. Xylem cells are tube shaped,
become hollow and die to form a pipe-like system through which water, hormones and mineral
nutrients move from the roots to the top of the tree.
Most of the radial growth of woody plants is due
to activity of the vascular cambium, but a small
amount results from activity in another lateral
meristem, the cork cambium, located outside the
vascular cambium. The cork cambium (phellogen)
together with the cork cells, constitute the periderm: a protective layer of suberized dead cork cells
forming the bark. Suberization is the impregnation
of cell walls of cork tissue with a fatty substance
called suberin. Each season new layers of cells are
produced and appear as growth rings when viewed
in cross-section. Over time, the xylem cells at the
center of the trunk or limb are crushed and become
nonfunctional as transport pipes, but they do provide structural support to hold the plant upright.
While grafting it is important to line up the cambiums of the scion and the rootstock to ensure a
successful graft union.

Buds
Buds are important to the vegetative and reproductive growth of trees. Fruit tree training and, to a lesser
extent, pruning primarily involves bud manipulation. Buds are actually undeveloped shoots. When
2

Classification by arrangement on the stem

The bud arrangement influences the arrangement
of a fruit trees branches and thus the trees shape
and how easy it is to manage. A node is the joint on
a stem where a leaf is or was attached (Figure 3).
Axillary buds are located in the axis above where a
leaf is attached to the stem. In apples there is usually only one leaf per node, whereas three leaves
often arise from a node on peach shoots. When a
leaf falls in the autumn, a leaf scar remains just
below the axillary bud (Figure 3). Buds are opposite when there are two at the same node but on
opposite sides of the stem. Forsythia is an example
of a plant with opposite buds. Buds are alternate
when there is only one from each node and no one
bud is on the same side of the stem as the one next
above or below it. Deciduous fruit trees have buds
that spiral along a shoot (Figure 4). The spiraling
three-dimensional arrangement of leaves around a
stem is known as Phyllotaxy and is expressed as
a fraction, where the numerator is the number of
turns to get to a leaf directly above another and the
denominator is the number of buds passed.

plant growth-regulator sprays, to manipulate tree

growth and flowering. Often these techniques
affect bud dormancy, so knowledge of buds and
bud dormancy is essential if we are to understand
how pruning influences tree growth. It is also
important to be able to identify the different types
of buds on a tree, especially to distinguish between
flower and vegetative buds.
Buds may be classified as to location, contents, or
activity.
Classification by content Several types of buds
commonly develop on fruit trees. Vegetative
buds only develop into leafy vegetative shoots.
Flower buds produce only flowers. Stone fruit
trees (peach, nectarine, apricot, plum, and cherry)
produce vegetative buds and flower buds. Apple
and pear trees produce vegetative and mixed
buds. Both leafy shoots and flowers emerge from
mixed buds.
Classification by location
Terminal buds are located at the tip of a shoot.
On stone fruit trees terminal buds are vegetative
buds. Terminal buds on apple and pear trees are
usually vegetative; however, some varieties such
as Rome Beauty produce mixed buds terminally
and are referred to as tip bearers or terminal
bearers. Most mixed buds on apple and pear
trees are formed terminally on short, less than sixinch, shoots that terminate with a rosette of leaves.
These short shoots are called spurs. Lateral buds
form in the axils of leaves and are often referred
to as lateral buds or axillary buds. On stonefruit trees, lateral buds may be either vegetative
or flower. Nodes on one-year-old shoots may have
one to three buds, some of which may be flower
buds and others vegetative buds. Flower buds are
larger with tips that are relatively round, whereas
vegetative buds are small, narrow, and pointed. In
the case of apple and pear trees, lateral buds on the
previous seasons growth are usually vegetative.
However, lateral buds on some varieties, especially on the dwarfing rootstocks, may be mixed
buds.

Terminal bud
Node
Internode
Node

Lateral bud

Leaf scar
Flower bud scar
Lenticel

Terminal bud scar

Figure 3. Section of a limb shows nodes, leaf scars, and different types of buds.

Plant Hormones

alternate
bud
arrangement

Hormones are substances produced in very small

amounts in one part of the plant and transported
to another part where they cause a response.
Plants produce a number of hormones that control
various aspects of growth, such as stem elongation; dormancy of buds and seeds; flowering;
fruit set, growth, and ripening; and the response
to light and gravity. While pruning, it is useful
to consider the activity of the general types of
hormones, promoters (gibberellins and cytokinins) and inhibitors (auxins and abscisic acid).
Promoters generally cause bud growth, cell division and elongation, and stem growth. Inhibitors
are usually associated with dormancy and inhibit
shoot development from seeds and buds and may
be involved in flower-bud induction. It is often
the ratio of promoters and inhibitors, rather than
their absolute concentrations, that determines
how a plant will grow. The production of plant
hormones is usually controlled by environmental
conditions such as temperature or day length.
Vegetative growth is usually associated with low
ratios of inhibitors to promoters and dormancy is
usually associated with high ratios of inhibitors to
promoters.

opposite
bud
arrangement

Figure 4. Shoots with alternate arrangement (left) and opposite arrangement (right). In each case the shoot has been
headed and the diagram to the right of the arrow indicates
how the buds respond to the heading cut.

Classification by activity
Buds are dormant when they are not visibly
growing. When shoots develop around large pruning cuts, they usually are sprouting from dormant
buds (Figure 5). Adventitious buds form irregularly on older portions of a plant and not at the stem
tips or in the leaf axils. They form on parts of the
root or stem that have no connection to the apical
meristems. They may originate from either deep
or peripheral tissues. For example, shoots often
arise from adventitious buds growing from callus
tissue around wounds. Root suckers (vigorous
upright shoots developing from the roots) develop
from adventitious buds on the roots.

Dormancy is a condition characterized by temporary growth cessation and suppressed metabolism. During the winter trees appear not to be
growing, but the tissues are alive, there is metabolic activity, and cells are slowly expanding and
differentiating. By early October all the flower
parts (petals, stigmas, anthers, etc.) can be seen in
a flower bud and vegetative buds contain leaves.
During the winter these various tissues continue
to enlarge and differentiate. Given favorable
growth conditions, some buds will develop into
shoots or flowers, but others may remain dormant. By understanding the factors influencing
bud dormancy we often can influence certain aspects of tree growth. Buds of deciduous trees go
through several stages of dormancy. Results from
dormancy research are confusing because plant
physiologists have used different terminology to
describe the stages of dormancy. Plant physiologists currently describe dormancy in four stages.

Watersprouts

Figure 5. Watersprouts developing from adventitious buds

around a pruning cut used to lower a tree.

Para-dormancy occurs in the mid to late summer

when buds do not grow because inhibitors produced in the leaves and terminal buds inhibit bud
growth. Para-dormancy can often be overcome by
removing leaves (leaf stripping) along a section of
a shoot so the axillary buds develop into shoots.
Nurserymen often use this technique to produce
trees with lateral branches (feathered trees). Using
heading cuts to remove the terminal portion of a
shoot will allow several axillary buds just below
the cut to develop into shoots. Sometimes an application of growth promoters (gibberellins and/or
cytokinins) will induce bud growth.
Sometimes axillary buds do not become dormant
and develop into shoots within a few days of being
formed. Such shoots are referred to as sylleptic
shoots and are fairly common on vigorously growing peach trees, but are rarely produced on apple
trees (Figure 6).
Ecto-dormancy occurs in the early fall, before
defoliation, when plants do not grow because
the environmental conditions are not conducive
for growth. Growth will resume if the plants are
exposed to suitable temperatures and day lengths.
Endo-dormancy occurs during the winter because
there are high levels of inhibitors (abscisic acid)
within the buds. During this phase of dormancy the

trees will not grow even under ideal growing conditions. The concentration of inhibitors declines
as buds are exposed to chilling temperatures.
Temperatures near 45F are ideal for chilling, but
temperatures between 35 and 55 F will provide
some chilling. The chilling requirement to satisfy
dormancy for most varieties of apples and peaches
grown in Virginia is about 1,000 and 800 hours,
respectively. When the chilling requirement is satisfied, the level of inhibitors within the bud is low
enough that growth may commence when environmental conditions are appropriate for growth.
Avoid planting varieties with chilling requirements
less than 800 hours because such varieties usually
bloom early and are susceptible to frost.
Eco-dormancy occurs in the late winter, usually
by mid January, after the chilling requirement has
been satisfied. At this time the trees do not grow
because conditions are unsuitable for growth.
Growth will commence when trees are exposed to
warm temperatures.
Apical dominance is a type of para-dormancy,
where axillary bud growth is inhibited in the api
cal meristematic zone. Axillary buds on fruit trees
typically remain dormant for a prolonged period
while the main shoot continues to grow. Apical
dominance has been studied for more than 80
sylleptic shoots

sylleptic shoots

Figure 6. Sylleptic-shoot growth on peaches during the growing season (left) and
during the winter (right). Arrows indicate sylleptic shoots.

Shoot bending
Shoots bend in response to an auxin gradient within
the shoot. Everyone who has grown plants in the
house has noticed that plants tend to grow towards
the light. This phenomenon is known as photomorphism and is caused by varying concentrations of
auxin in different sides of a stem or shoot. Auxin
causes cells to elongate, but auxin is destroyed by
light. Therefore, there is a higher concentration
of auxin on the dark side of a shoot and the cells
on the dark side elongate more than cells on the
sunny side of the shoot, causing the shoot to bend
towards the light (Figure 8).

years, and the exact mechanism is not yet fully

understood, but it seems to be controlled by the
relative concentrations of inhibitors and promoters. Growth of axillary buds is inhibited by high
concentrations of auxin produced by the terminal
bud. Auxin moves down the shoot, from cell to cell
by gravity, so concentrations are highest near the
shoot tip. Promoters are produced in the roots and
are transported upward in the tree. Growth of axillary buds may occur at the base of shoots where
concentrations of inhibitors are relatively low and
concentrations of promoters are relatively high.
You can overcome apical dominance by removing
the shoot tip, which is the source of auxin (Figure
7). The three or four buds immediately below a
heading cut usually develop into shoots. Pinching
annual plants to induce branching is a form of heading. Another way to overcome apical dominance is
to notch buds. Notching involves cutting through
the bark to hard wood, with a knife or hacksaw
blade at about bloom time, just above a bud. The
cut interrupts the downward flow of inhibitors, but
not the upward flow of promoters, and releases
the bud from dormancy. On vigorous upright oneyear-old shoots, notching often successfully overcomes dormancy in about 70 percent of the buds.
Sometimes apical dominance can be overcome by
spraying shoots with promoters (gibberellins and/
or cytokinins) just before bloom time.

Unpruned
B

Figure 8. Photomorphism is the bending of a shoot towards

the light. The auxin concentration is highest on the dark
side of the stem and causes cells on that side to elongate,
resulting in stem curvature.

Figure 7. One way to overcome apical dominance and

Dormant
A

Light

Light

Tree fruit producers have noticed a similar phenomenon where the tips of growing branches
tend to bend upward, even when the branch was
physically oriented to the horizontal. This condition, known as gravimorphism, is also caused by
an auxin gradient within the branch in response to
gravity. Auxin flows by gravity to the lower side
of a limb. The subsequent accumulation of auxin
is responsible for increased cell elongation on the
underside of the limb, and the growing tip bends
upward.
Another consequence of gravimorphism is the
development of watersprouts from the upper surface of horizontally oriented limbs. Watersprouts
are vertically growing shoots that develop from the
upper surfaces of branches or near pruning cuts.

Heading
cut
C

inducing branching where we want branching is to head the

shoot (A). If the shoot is not headed, the top several buds
will develop into shoots (B). If the shoot is headed, several
buds below the heading cut will develop into shoots (C).

Pruning reduces yield

Pruning removes wood with flower buds, and thus
potential fruit. Yield from pruned trees is nearly
always less than yield from nonpruned trees, but
fruit quality is improved by pruning. Pruning
improves fruit size by increasing the amount of
leaf area per fruit. Pruning improves light distribution throughout the tree, which is important for the
development of fruit red color and sugar levels.

High auxin concentrations on the underside of the

limb inhibits growth of buds on the underside of the
limb, but the concentration of auxin on the upper
side of the limb is inadequate to inhibit bud growth
and many of these buds develop into watersprouts.
Watersprouts are usually undesirable and their
development can be suppressed by orienting limbs
no more than 45 degrees from the vertical. Fruit
trees are sometimes trained as espalier (tree fence).
There are several ways to espalier trees, but one
method involves orienting limbs to a horizontal
position. This system induces many watersprouts
along the length of the branches. Watersprout
development can be greatly suppressed by orienting
limbs 45 to 60 degrees above horizontal (Figure 9).

Pruning delays fruiting

Pruning encourages vegetative growth rather than
reproductive growth in young trees. A nonpruned
tree will always flower and produce fruit earlier
in the life of the tree than a pruned tree. The reason young trees are pruned is to induce branches
to develop where they are wanted and to develop
a strong tree structure that will support large crops
as the tree matures. As a tree matures the physiology changes from vegetative growth to reproductive growth. To obtain high annual yields of mature
trees, it is important to minimize fruiting until trees
have nearly filled their space. Pruning is one technique used to delay fruiting of young trees.

Reducing tree height by cutting into large diameter branches or trunks often results in the development of vigorous watersprouts around the cut.
There are buds buried in the bark that normally
remain dormant. However, a severe pruning cut
will release these buds from dormancy.
40

60

70

80

Summer pruning

Summer pruning involves the selective removal of

leafy shoots during the growing season. Responses
to summer pruning vary with time of pruning,
severity of pruning, tree vigor, geographical location, and variety. Several researchers evaluated
summer pruning during the 1980s and several general statements can be made about the practice.
Summer pruning reduces within-tree shade and
usually improves fruit red color development and
sometimes improves flower bud development.
Summer pruning removes leaves that produce photosynthates (sugars) for growth of all tree parts.
Summer pruning sometimes reduces fruit size and
sugar levels.
Due to reduced whole-tree photosynthesis, summer pruning suppresses late-season trunk enlargement and root growth.
Summer pruning does not suppress shoot elongation the following season. Summer pruning
reduces late-season photosynthesis, and theoreti-

C
D

Figure 9. Auxin distribution within a stem is controlled by

gravity. When limbs are oriented from vertical to about
60 degrees from vertical, auxin is distributed fairly evenly
around the limb and buds develop into shoots fairly symmetrically around the limb (A and B). Auxin accumulates
on the underside of flat limbs (C and D) and inhibits growth
of buds on the underside. Auxin concentration is low on
the upper side and buds are not inhibited and develop into
strong watersprouts.

Additional Pruning Facts

Pruning is a dwarfing process
Pruning increases vegetative growth near the
pruning cut and this gives the illusion that pruning
stimulates growth. However, the weight of a tree
that was pruned annually is always less than the
weight of a nonpruned tree.

cally should reduce the accumulation of reserve

carbohydrates within the tree that are used for
early season growth. However, results from most
pruning experiments indicate that the response to a
certain type of pruning cut will be the same regardless of the time of year the cut was made.

Training and Pruning Apples Trees, Virginia Cooperative Extension publication 422-021, http://pubs.
ext.vt.edu/422-021/
Training and Pruning Apple Trees in Intensive
Orchards, Virginia Cooperative Extension publication 422-024, http://pubs.ext.vt.edu/422-024/
Pruning Peach Trees, Virginia Cooperative
Extension publication 422-020, http://pubs.ext.
vt.edu/422-020/

Summary

Pruning is an important orchard practice because

pruning can influence fruit quality and the balance
between vegetative growth and fruiting. Successful pruners observe how plants respond to various
types of plant manipulation, including pruning.
Profitable fruit production requires an understanding of plant physiology, and how pruning alters the
physiology of the plant. For further information
concerning how to prune fruit trees, see the Extension publications that provide information on how
to prune and train apple and peach trees.

Reviewed by Tony Wolf, Extension specialist, Alson H.

Smith, Jr. Agricultural Research and Extension Center