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14 Pruning and Training Physiology

David C. Ferree1 and James R. Schupp2


1Department of Horticulture and Crop Science, Ohio Agricultural Research and
Development Center, Ohio State University, Wooster, Ohio, USA;
2Department of Horticultural Sciences, Cornell University, Highland, New York, USA

14.1 Introduction 320


14.2 Pruning 320
14.2.1 Effects on growth 320
14.2.2 Effects on flowering 322
14.2.3 Effects on fruit 322
14.2.4 Effects on mineral nutrients 322
14.3 Types of Pruning Cuts 322
14.3.1 Thinning cuts 323
14.3.2 Heading cuts 324
14.4 Time of Pruning 324
14.5 Pruning Trees of Different Ages 325
14.5.1 Young trees 325
14.5.2 Fruiting trees 327
14.5.3 Old or declining trees 330
14.6 Summer Pruning 331
14.6.1 Effects on physiological processes 331
14.6.2 Effects on fruits 331
14.6.3 Effects on growth 333
14.6.4 Practical implications 333
14.7 Root Pruning 334
14.7.1 Effects on physiological processes 334
14.7.2 Effects on growth 335
14.7.3 Effects on fruiting 336
14.7.4 Practical implications 336
14.8 Training 337
14.8.1 Bending 337
14.8.2 Phloem interruption 339
14.8.3 Notching 341
14.9 Conclusion 341

© CAB International 2003. Apples: Botany, Production and Uses


(eds D.C. Ferree and I.J. Warrington) 319
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320 D.C. Ferree and J.R. Schupp

14.1 Introduction of this chapter is to provide the physiological


basis for these cultural techniques so that
Pruning and training are almost always they can be used to best advantage.
mentioned together, but there is a difference
between them and each has a clear function.
Pruning has been defined as the art and 14.2 Pruning
science of cutting away a portion of the plant
for horticultural purposes. Pruning can be 14.2.1 Effects on growth
used to improve tree shape, to influence its
growth, flowering and fruitfulness, to improve Although the practice of pruning goes back
fruit quality, to repair injury, to contain the several thousand years, the science of prun-
plant and to encourage light and spray ing originated in the early 20th century, with
penetration. Training, in contrast, refers to experimental data revealing several general
the direction of tree growth or form and the influences. The first of these is that pruning
development of the structural framework of is a dwarfing process and the more severely
the tree. Pruning is only one of the tech- a plant is pruned, the greater the degree of
niques used in training. Although some dwarfing. Removal of the branch not only
training may be necessary after the tree removes stored carbohydrate and nitrogen
comes into production, training is mostly reserves, but also reduces potential leaf sur-
confined to the period when the tree is face and growing points as well. Obvious
becoming established. The ultimate goal of reductions in number of growing points, tree
both pruning and training is to improve light height and spread and leaf area can easily be
distribution so that as much of the tree observed on pruned versus non-pruned
canopy as possible maintains production of plants. Although not as obvious, pruning
high-quality fruit. Forshey et al. (1992) pre- correspondingly reduces root growth, and
sent a comprehensive treatise on pruning new root growth will be delayed until shoot
and training, including an extensive refer- growth in response to pruning occurs.
ence list. A comprehensive literature review There is a fundamental equilibrium
will not be included in this chapter but refer- between the above- and below-ground
ences will be cited to illustrate key points. components of an apple tree. Removal of a
Apple trees that are not trained and portion of either the top or root system slows
pruned become very large (Plate 14.1) and growth of the other until the balance is re-
develop fine branching at the tree periphery, established (Taylor and Ferree, 1981; Geisler
which becomes very dense and shades the and Ferree, 1984a). This equilibrium is proba-
tree interior. Fruit size, colour and sugar con- bly achieved not only through loss of reserves
centration decline on shaded spurs and, as in the tissue removed, but also through
light becomes still more limiting, the spurs changes in the functional equilibrium
themselves become weak and unproductive, between the root and the top and the balance
eventually dying. As the trees age, a larger of hormones produced in meristematic
and larger portion of the tree canopy volume regions in the leaves, shoot tips and root tips.
becomes unproductive. As the unproductive Shoot growth is dependent upon water
area of the canopy expands, it becomes a and mineral nutrient uptake by the root sys-
larger sink for photosynthates and thus tem, while roots depend upon the above-
decreases tree efficiency. The dense shading ground portion of the tree for carbohydrates.
encourages insects and diseases due to high Pruning away a portion of either of these
humidity, slow drying conditions and poor plant parts reduces its capacity to perform its
spray penetration. Thus, although pruning function; thus growth in other parts of the
and training are expensive cultural practices, plant is reduced and energy is redirected to
economic fruit production is not possible regenerate the missing component. This con-
without the improvement in light distribu- cept, known as functional equilibrium,
tion these techniques provide to maintain seems to be moderated by plant hormones
tree efficiency and fruit quality. The purpose (Richards and Rowe, 1977).
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Pruning and Training Physiology 321

Grochowska et al. (1984) have shown that work of Verner (1955) illustrated that when
pruning the top during the dormant season four or more shoots were above a new
increases the supply of cytokinins from the branch, the angle of that new branch was
roots, measured as increased concentration in 50–90° (i.e. close to horizontal). When there
the remaining above-ground tissue. These were no shoots above the new branch, the
increased hormone levels are probably respon- branch angle was in the very narrow range
sible for stimulating cell division and ulti- of 0–40° (i.e. very upright). Removal of the
mately shoot growth, which, in turn, promotes terminal bud results in an increase in the
auxin production in the shoot tips and gib- growth of the subtending laterals, and
berellin production in the new, unfolding shoots that would have been spurs become
leaves. The increases in concentration of these vegetative shoots. Removal of apical domi-
three classes of hormones found in shoot tissue nance not only changes which shoots grow
in the early part of the growing season follow- and their vigour, but the resulting shoots are
ing dormant pruning not only increase growth more upright with narrower crotch angles.
but also alter the pattern of growth. Removal Thus, tree form is changed into a more
of terminal buds and their auxin results in upright, narrow shape.
enhanced growth of laterals of latent and Even though pruning dwarfs the whole
existing flower buds. The more upright the plant or limb, an invigorating growth
branch that is pruned, the more vigorous will response occurs in the immediate area of the
be the growth that responds to the removal of cut. This invigorating effect decreases as the
apical dominance (Myers and Ferree, 1983b). distance from the cut increases. This is an
Normally a terminal bud makes the most important general concept to understand,
growth in an unpruned, upright branch, and because, if the purpose of pruning is to
each subsequent subtending lateral makes invigorate and stimulate growth, as is often
progressively less growth, until some form the case in areas of the canopy that are spur-
spurs, with the most basal buds not grow- bound, more response is generated by many
ing. This growth form is probably the result small cuts than by a few large ones (Ferree
of cumulative auxin concentrations pro- and Forshey, 1988). Generally, many small
duced from each of the buds above causing cuts stimulate more shoot growth, reduce
a reduction in growth in the shoot originat- fruiting to a greater degree and alter tree
ing at lower levels (Fig. 14.1). The classic form more than when a few large cuts are
made. Larger cuts result in fewer, more vig-
orous and more localized regrowth. Another
manifestation of the localized influence can
be observed in the injury that occurs when
trees are pruned close to a sudden drop in
winter temperature. The freezing injury
results in death of the bark and phloem
tissue and is most obvious close to pruning
cuts, at the tree base and in limb crotches
on pruned trees compared with unpruned
adjacent trees (Rollins et al., 1962; D.C.
Pruning cut Ferree, personal observation). This localized
response occurs even though the tree appears
to be completely dormant.
One of the primary functions of pruning
is to develop a framework of branches that
are strong, likely to consistently carry a
good crop of premium-quality fruit and be
Fig. 14.1. Natural growth pattern of apple (left) and less prone to environmental extremes. To
change in the pattern caused by a heading cut achieve this goal the crotch angle, which
(right). forms very early in the first year of the
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322 D.C. Ferree and J.R. Schupp

development of a branch, must be wide and pruned trees because the supply of reserves
strong. Narrow crotches cause bark inclu- available to remaining blossoms is
sions, are weak, split easily and are more sus- increased. Cell division early in the season is
ceptible to cold injury. Wide crotch angles are greater in areas of the canopy that have high
formed naturally in very vigorous-growing light and large spur-leaf areas. Later in the
trees that have very active meristematic season during cell expansion, shoot leaves
areas that produce large quantities of auxin supply the fruit. Fruit size is closely related
(Verner, 1955). to spur-leaf area and well-illuminated
shoots (Ferree and Palmer, 1982; Rom and
Ferree, 1986). Pruning is the cultural practice
14.2.2 Effects on flowering most influential in establishing a canopy
light environment that promotes high spur
Another general principle is that pruning quality and an even distribution of shoots
delays the flowering and fruit bearing of through the canopy.
young trees and decreases fruiting in
mature trees. Forshey et al. (1992) state that
this is the result of the following three fac- 14.2.4 Effects on mineral nutrients
tors: (i) the removal of effective bearing sur-
face; (ii) the stimulation of vegetative Another indirect effect of pruning is the
growth from growing points that were for- influence of pruning on the mineral content
mer or potential fruiting spurs; and (iii) of the tree and fruit. The decrease in number
direct competition between vigorous shoot of fruits caused by pruning is associated
growth and fruit set. They indicate that the with an increase in leaf nitrogen, potassium
reduction in cropping is proportionately and phosphorus and a reduction in fruit
much greater than the reduction in total concentrations of magnesium and calcium.
growth. Pruning young trees tends to force These changes are associated with the com-
growth of long succulent shoots, which con- petition of the fruits and vigorous shoots for
tinue to grow late in the season. Thus, car- the elements and with the increase in fruit
bohydrates do not accumulate for size caused by dormant pruning. Since cal-
flower-bud initiation, but are used in pro- cium is so important to fruit storage poten-
ducing the vigorous vegetative growth tial and disorders such as bitter pit, the
(Mika, 1975). Elfving (1990) demonstrated reduction in fruit calcium caused by pruning
that heading back pruning in a single year is a particular concern. Since vigorous shoots
significantly reduced tree size and yield and are a stronger sink for calcium than the fruit,
the effect remained evident 5 years after the dormant pruning reduces fruit calcium and
pruning. The more severe the pruning, the the more severe the pruning, the greater the
greater is the decrease in fruiting (Elfving reduction. Dormant pruning also increases
and Forshey, 1976). The negative influence fruit size, which leads to a dilution effect on
of pruning on fruiting is particularly impor- calcium concentration. In contrast, summer
tant in intensively planted modern orchards pruning, which removes vegetative shoots,
that are dependent on early cropping to can increase fruit calcium (Preston and
help control growth. Perrin, 1974).

14.2.3 Effects on fruit 14.3 Types of Pruning Cuts

One of the primary reasons for annual prun- All pruning cuts can be classed into two
ing is to increase fruit size. The effect on types (thinning out and heading back) and
fruit size is not a direct effect of pruning, but each type of cut produces a different physio-
results from the reduced number of fruit logical response. A thinning cut (Fig. 14.2)
and improved light conditions in the removes an entire shoot, spur, branch or
canopy. Generally fruit set is higher on limb, while a heading cut (Fig. 14.3) removes
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Pruning and Training Physiology 323

Dormant Unpruned Pruned – thinning cuts


Fig. 14.2. Change in growth of an apple tree that received two dormant thinning cuts in year 1 (left-hand
figure). Growth response at the end of year 2 is shown in the right-hand figure. The response for an
unpruned tree is shown in the centre figure. In the latter case, main pruning cuts are needed in the second
dormant season to restore a desired tree form but note that the advantages of lateral branch development
and uniform fruit distribution have been lost.

Dormant Unpruned Pruned – heading cuts


Fig. 14.3. Change in growth of an apple tree that received three heading-back pruning cuts in year 1
(left-hand figure). Growth response at the end of year 2 is shown in the right-hand figure where lateral
branch development is strong and vigorous upright growth is apparent. The unpruned tree (centre figure) has
a more spreading growth form with less lateral branching.

only a portion of a shoot or limb, leaving remove unfruitful wood that is upright and
another portion from which new growth can vegetative or pendant and weak.
develop. A comparison of the various Generally, thinning cuts improve light dis-
responses from the two types of cuts is sum- tribution in the tree, thus increasing flower
marized in Table 14.1. initiation, fruit set and spur growth in lower
parts of the canopy. The effect of thinning cuts
on fruiting is limited to the amount of poten-
14.3.1 Thinning cuts tial bearing surface removed. The ratio of ter-
minal to lateral buds is largely undisturbed
Thinning cuts are primarily used to alleviate and, by removing branches at their point of
crowding and shading and generally origin, only weaker latent buds are left to
improve light penetration into the canopy grow; thus, thinning out does not increase
(Fig. 14.2). Thinning cuts also function to shoot growth as much as heading back.
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324 D.C. Ferree and J.R. Schupp

Table 14.1. Relative influence of thinning and heading pruning cuts on apple trees.

Response Thinning cuts Heading cuts

Apical dominance Little effect on remaining growth Terminal removed, buds released
Shoot formation Fewer vegetative shoots Increased number and length
Spur formation Increased Decreased
Tree shape Limited effect Dramatic effect, e.g. hedging
Tree growth habit Natural tall rangy Compact and upright habit
Fruitfulness Limited effect Significant decrease
Carbohydrate reserves Limited effect Dramatic reduction
Nutrition elements Limited effect Decrease calcium, magnesium,
increase nitrogen
Local stimulus Encourage few shoots Encourage many shoots

The thinning-out cut should be just above growth and fruiting lasting 5 years.
the small ridge at the base of a branch, called Heading cuts stiffen limbs and care should
the collar. Cuts made at this location make be exercised in the use of these cuts on
the smallest wound, heal the fastest and are upright branches of young trees because
less susceptible to winter injury. Leaving a some type of physical limb bending will be
stub above the ridge or cutting too close and required to return these branches to a bal-
removing the ridge slows healing and pro- anced fruiting condition.
vides more potential for disease or insect Each type of cut has a role in creating and
damage (Shigo, 1990). An exception to this managing an efficient tree with a good bal-
rule is the Dutch cut, a bevelled stub that is ance between vegetative vigour and fruiting.
left for stimulating a renewal limb with cer- Newly planted trees are headed to stimulate
tain training systems (see Chapter 15). the production of strong lateral branches
needed to form scaffolds. Thinning cuts are
necessary on young trees to develop the
14.3.2 Heading cuts desired primary branch framework with
minimal effect on fruiting. As trees reach full
Heading cuts (Fig. 14.3) locally stimulate size, thinning cuts maintain the adequate
more vegetative growth, which tends to be light and spray penetration needed to keep
vigorous and shades the tree interior, result- the whole canopy productive, while fruit
ing in fewer fruiting sites and less flower size and quality are improved. Cultivars
initiation than thinning cuts. Since using this such as spur-type ‘Delicious’ often form
type of cut removes many terminal buds, insufficient numbers of laterals and may
apical dominance and the hormone balance benefit from heading cuts to induce laterals
is disturbed more than with thinning cuts. where they are needed. In older trees, when
Heading increases both shoot number and portions of the canopy become weak or spur-
length, but the effect on length is determi- bound, heading cuts can encourage shoot
nate. The younger the wood and the more growth and improve fruit size.
upright the growth headed, the greater the
effect. Heading 1-year-old wood results in a
strong localized response, releasing the 14.4 Time of Pruning
three to four buds below the cut to develop
into vigorous, upright shoots with narrow Most pruning on apple trees is done when
crotches. Heading causes many shoots that they are dormant. In areas that can experi-
would have been potential spurs or fruiting ence very low temperatures in winter, time
shoots to form vigorous vegetative growth. of pruning is an important consideration
Elfving (1990) has shown that heading cuts because it increases susceptibility to low-
on young trees have negative effects on temperature injury (winter injury). Sensi-
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Pruning and Training Physiology 325

tivity to cold injury after pruning is greatest 14.5 Pruning Trees of Different Ages
in the tissue in close proximity to the prun-
ing cuts. The exact temperature required to 14.5.1 Young trees
induce winter injury varies with many fac-
tors, such as previous exposure to low tem- Most apple orchards are trained to some
peratures, previous crop, cultivar, rootstock modified form of a central leader system,
and tree age. Winter injury may occur at resulting in a cone-shaped canopy. In this
low temperatures in the range of 23 tree form, the central stem, or leader, is
to 49°C, depending upon the inherent maintained in an upright orientation, while
hardiness of the tree and these contributing limbs that form on the sides are trained at an
factors. Pruning within 2 weeks prior to the angle to form the primary scaffolds.
low-temperature event will increase the Additional secondary limbs form on both the
sensitivity of the tree to cold injury, with the leader and the scaffolds and are maintained
greatest loss of hardiness occurring within to provide a bearing surface. The purpose of
the first 48 h (Rollins et al., 1962). The loss in this chapter is not to provide a ‘how-to’ trea-
hardiness is then gradually regained. Thus, tise on pruning trees trained to specific sys-
the risk of causing winter injury by pruning tems, but to cover the principles and
can be reduced by suspending pruning physiological basis for practical application
work when severe cold weather is forecast for a generalized central-leader system. The
(see Chapter 10). specific alternative pruning practices for
To reduce the risk further in areas prone young trees in specific training systems will
to winter injury, pruning should be delayed, be covered in Chapter 15.
where practical, until after the coldest months Trees that come from the nursery have
(January and February in the northern lost a significant amount of their root sys-
hemisphere). If pruning must be started ear- tem and thus the top needs to be reduced
lier, it is best to prune the oldest trees and to create a balance so that sufficient growth
most cold-tolerant cultivars first and to delay occurs to form the permanent scaffolds. If
pruning the youngest trees and most cold- trees arrive as unbranched whips, they are
sensitive cultivars. usually headed back 60–90 cm above the
When pruning is delayed until growth ground, depending on which training sys-
starts, the tree responds differently. Delay tem is used. Cultivars such as spur-type
much beyond bloom devitalizes growth and ‘Delicious’ do not branch easily and will
may interfere with the development of only produce shoots within 5 cm of the
flower buds for the next year’s crop. Pruning heading cut, while cultivars such as
cuts at the time vegetative growth is begin- ‘Golden Delicious’ will produce more later-
ning result in an increase in the number of als, which are distributed 10–15 cm below
buds that break and buds further from the the cut (Ferree, 1978). Cultivars that have a
cut tend to break more when compared with terminal bearing habit should be headed
dormant pruning. This response is probably high, as these cultivars tend to produce
due to the removal of apical dominance since weaker shoots from the lower buds, plus
the meristematic regions in the growing the terminal bearing habit causes the
buds are primary sources of auxin. Generally branches to bend, which can cause them to
the breaks that occur from delayed pruning interfere with herbicide applications and
are less vigorous and may be very desirable mowing operations if they originate too
on cultivars that tend to have blind wood low on the trunk.
(previous-season wood with no lateral If feathered trees (see Chapter 6) are
growth), such as ‘Tydeman’s Red’ or ‘Rome planted, shoots to form the primary scaffolds
Beauty’. Delaying dormant pruning until are already in place and, generally, the only
bloom can also be used to lessen the cuts made on these trees are to remove any
regrowth in overly vigorous trees; however, spurs or shoots below 45 cm above the
delayed pruning should not be used on trees ground, as these interfere with weed control
with less than optimum vigour. and will produce few marketable fruit.
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326 D.C. Ferree and J.R. Schupp

During the first growing season, leader of the remaining leader shoot. Cultivars such
and scaffold selection are very important as spur-type ‘Delicious’ may not produce
and should be done early so that the tree sufficient shoots to use this method of leader
directs its growth as much as possible into selection. In this instance, the shoots with
tissues that are to be retained. Each growing upright growth and very narrow crotch
shoot produces auxin, which moves down angles, which would be removed in the
the tree and affects the growth and crotch above procedure, are retained. A spring
angle of shoots below. The uppermost shoots clothes-peg is clamped around the leader
grow vertically because of an absence of just above the young shoot when it is 5 cm
auxin during and after bud break. The pro- long, and it is forced to grow horizontal
gressively increasing auxin concentration below the clothes-peg (Fig. 14.4). This shoot
further down the trunk stimulates lower will ultimately curve upward, but only after
shoots to develop wider crotch angles and a wide, strong, crotch angle has been estab-
this effect is complete by the time the shoots lished. The growth of these shoots is reduced
are 5–10 cm long (Verner, 1955). This natural 15–20% by the bending process further
pattern can be used to advantage by select- establishing the dominance of the leader. An
ing the leader when the shoots are 10 cm alternative to the clothes-pegs is to use
long, removing the two or three upright pointed toothpicks or thin wire to force the
shoots that are growing just below the leader new shoot to grow laterally. These practices
and leaving the lower shoots with wider must be carried out before the base of the
crotch angles to develop as scaffolds. Since shoot lignifies. It takes about 1 min per tree
the tissues in question are succulent, care to install the clothes-pegs and about 15 s to
must be exercised to avoid injury to the vas- remove them 3–4 weeks later to avoid
cular tissue, as this can decrease the growth girdling the leader.

Headed, Headed, Headed,


untreated clothes-pegs in place subsequent form following
use of clothes-pegs

Fig. 14.4. Influence of clothes-peg spreading on growth of a newly planted unbranched apple tree that has
been headed.
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Pruning and Training Physiology 327

The most important principle to remem- Vigour varies throughout the canopy, with
ber on young trees is to prune them as little greatest vigour in the top and periphery. This
as possible. The delay in cropping caused by variability is mostly due to light exposure and
heavy pruning has an adverse effect on one of the main goals of pruning is to create
early profitability. In intensive plantings the the optimum balance of vigour and fruiting in
vigorous growth caused by heavy pruning as much of the canopy as possible by improv-
may negate much of the advantage in early ing canopy light distribution. The orientation
yields that are achieved by using increased of fruiting branches affects vigour and relative
tree numbers. response to pruning (Fig. 14.5). Normally
upright branches are mostly vegetative and
respond to pruning by growing vigorously. It
14.5.2 Fruiting trees is best to remove these by thinning cuts to
avoid shading portions of the canopy below
Although young trees are often handled them. Pendant branches are often weak and
similarly, as the tree begins to fruit, pruning have low fruit set and poor fruit size, colour
practices must be adjusted to the vigour of and quality (Tustin et al., 1988). These branches
the cultivar and rootstock (see Chapter 5) should be removed to eliminate the small,
and the growth and fruiting habit of the cul- low-quality fruit they produce. Branches with
tivar. Tree vigour is influenced by many fac- an orientation between horizontal and 45º
tors. Fertile, well-drained soil results in above horizontal are the most fruitful, have
much greater vigour than shallow, poorly the highest fruit quality and should form the
drained soil. Some climates are ideal and greatest portion of the fruiting canopy. Even
promote vigorous growth, while others distribution of light across these branches is
cause stresses that reduce growth. Cultural maintained by thinning-out cuts.
practices, such as nitrogen fertilization and Cultivars respond differently to pruning
weed and soil management, influence and training due to differences in their overall
growth. All these factors interact with the level of vigour, growth and fruiting
genetic potential of the scion and rootstock (Table 14.2). Growth habit refers to the overall
and must be considered when judging the growth pattern of the tree, including the
responses of the fruiting tree to pruning. degree of branching, the branch orientation

Fig. 14.5. Influence of branch orientation on growth and fruiting of apple.


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328 D.C. Ferree and J.R. Schupp

Table 14.2. Relative vigour of some commercially important apple cultivars and their growth and fruiting
habit according to Lespinasse (1980).

Low vigour Medium vigour High vigour

‘Braeburn’ (II) ‘Cox’s Orange Pippin’ (III) ‘Cortland’ (IV)


‘Idared’ (II) ‘Delicious’ (II) ‘Granny Smith’ (IV)
‘Jonathan’ (III) ‘Elstar’ (II–III) ‘Gravenstein’
‘Rome Beauty’ (IV) ‘Empire’ (I) ‘McIntosh’ (III)
‘Spur Delicious’ (I) ‘Fuji’ (IV) ‘Mutsu’ (III)
‘Spur Golden Delicious’ (I) ‘Gala’ (III) ‘Northern Spy’ (II)
‘Spur McIntosh’ (I) ‘Golden Delicious’ (III) ‘Spartan’ (II)
‘Tydeman’s Red’ (IV) ‘Jonagold’ (II)

(upright or spreading) and the branch crotch Type II is characterized by standard-habit


angle. Fruiting habit refers to the overall pat- ‘Delicious’. Branching is more frequent than
tern of fruiting and includes fruit position on in Type I and there is a tendency for the
the ends of long or short shoots, age of spurs fruiting zone to move away from the trunk.
producing most of the crop and location of the The trees have a tendency to develop narrow
crop on the scaffold limbs. Lespinasse (1980) crotch angles and benefit from spreading.
developed a system of classification for Type II trees may develop many medium-
growth habits (Fig. 14.6) and these have strong sized branches, resulting in a too dense
implications for the type of pruning required. canopy if not thinned out. No more than
Spur types, characterized by spur strains four permanent branches per tier should be
of ‘Delicious’, are classified as Type I. These retained. More thinning cuts into younger
tend to form few laterals on main scaffold wood are required to induce spurs and retain
limbs and have strong basitonic characteris- the fruiting zone in the tree interior.
tics (the most vigorous growth is at the base Type III cultivars are characterized by
of the tree). Thus the pattern of growth in ‘Golden Delicious’ and tend to be spreading
type I cultivars is characterized by strong with wide crotch angles and frequent
scaffold limbs breaking at the base of the branching. Cultivars of this type tend to bear
tree. The dominance of the central leader is early, with most of the fruit on spurs or
lost quickly unless shoots intended as scaf- terminals of short shoots borne on 2- to
folds are mechanically spread to weaken 4-year-old wood. The fruiting zone on
their growth relative to that of the leader. cultivars in this classification tends to move
Heading cuts may be needed to develop lat- away from the trunk to the outside of the
eral branches on primary scaffolds, which canopy. This requires thinning on the canopy
forces would-be spurs to grow into vegeta- periphery and heading cuts to lateral
tive extension shoots. Due to the sparse branches on the bases of scaffolds to induce
branching habit more scaffolds may be left, renewal of fruiting wood. Trees of this type
but care must be taken that they are spread are the easiest to adapt to a wide range of
vertically along the leader or the dominance training systems and are generally the easiest
of the leader and its growth will be reduced. to manage in the orchard.
Fruiting occurs on numerous short spurs, Type IV, the tip bearers, are characterized
which are long-lived. The zone of fruiting by ‘Rome Beauty’ and ‘Granny Smith’. They
remains close to the trunk and to the base of tend to have upright main scaffold limbs
the scaffolds as long as sufficient light is with narrow crotches and frequent branch-
maintained in these canopy areas. Type I cul- ing. A weeping terminal habit develops since
tivars tend to be prone to biennial bearing most of the crop is produced on the ends of
and can easily become spur-bound and the previous year’s shoots. The lower half of
senescent, particularly if they are propagated many shoots will be devoid of leaves or fruit
on very dwarfing rootstocks. on young trees, a condition known as ‘blind
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Pruning and Training Physiology 329

Fig. 14.6. Lespinasse (1980) classification of apple-cultivar growth habits.

wood’. Delayed spring pruning can often as ‘Empire’, require special treatment during
induce laterals in this branch region, which the developmental years if they are to be suc-
would normally be bare. This growth habit cessfully grown and managed in training
tends to be the most annual in production and systems that depend on a defined leader.
is often associated with high productivity In a young fruiting orchard, leader man-
and large fruit size. agement and, ultimately, tree height require
Low-vigour cultivar-and-rootstock com- consideration. As planting density intensi-
binations can be pruned harder than high- fies, it is critical that one row of trees does
vigour combinations without upsetting the not shade the adjoining row. Several studies
balance of growth and fruiting. Some root- show that a north/south orientation results
stocks, such as Oregon 1, cause upright in more even canopy light distribution. In
growth, while others, such as M.9 or M.9 model studies (Cain, 1972; Palmer, 1989) it
interstems, cause an open-spreading canopy. has been shown that tree height should not
Rootstocks that promote an open-spreading exceed twice the clear alley width. If a 2.5 m
canopy combined with cultivars with little free alley is needed for spray equipment and
tendency to maintain a central leader, such removal of fruit at harvest, then trees should
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330 D.C. Ferree and J.R. Schupp

not exceed 5 m in height. If this is exceeded, fied on young fruiting trees and the indis-
much of the row will be shaded in the after- criminate cuts from mechanical hedging gen-
noon by neighbouring rows. erally result in uneconomic production.
In-row spacing also has an impact on
leader management. Tustin et al. (1998) have
shown that creating rows with a sawtooth 14.5.3 Old or declining trees
profile from creating a gap between leaders
of adjoining trees results in improved light As trees age, fruit size generally declines and
distribution and improved fruit quality. To it requires a more detailed and time-consum-
achieve these goals there must be strong ing pruning job to achieve adequate fruit
vigour in the leader when the trees are size and quality. Trees can become spur-
young. This is achieved by removing shoots bound and perform as old trees even though
competing with the leader. In cultivars such they are not old (Ferree and Forshey, 1988).
as ‘Empire’ that fail to develop strong lead- These trees often have very large spur com-
ers, the leader should be headed by a third plexes with many weak growing points and
each year and then singled to one shoot early have very little shoot growth in the lower
in the growing season. When the tree reaches portion of the canopy. Spur pruning that
the desired height, the leader can be cut back removes 30% of the growing points on each
to an upward-growing shoot each year, just spur complex, plus a series of heading cuts
below the desired height, and the competing over the canopy can improve fruit size in
shoots removed. In supported systems it is these trees.
possible to let the leader fruit and bend over Another problem common in older
and after fruiting cut to one upright shoot orchards is that too many limbs have been
and repeat this process every couple of allowed to develop in the top of the trees.
years. Handling tree height in systems such The upper limbs in the tree shade the lower
as the slender spindle, which requires a limbs, and this uneven light distribution lim-
weak leader, or systems that have more than its production and fruit size in the bottom
one leader is covered in Chapter 15. half of the canopy. Often the productivity
Some very large orchards attempt to and profitability of old trees are limited by
reduce the cost of pruning by mechanically being both top-heavy and spur-bound.
hedging or pruning only every second or A comparison of several corrective prun-
third year. Mechanical pruning unfortu- ing strategies for top-heavy, spur-bound
nately causes mostly heading cuts, and the ‘Starkrimson Delicious’ trees showed that
vigorous upright shoot growth that follows removing several large limbs in the top of
quickly produces heavy shade in the tree, the canopy or making numerous heading
which reduces flower formation and fruit cuts improved light penetration, improved
quality. This effect is particularly bad if fol- fruit size and increased fruit quality, but the
low-up manual pruning is not used to thin reduction in yield in these treatments
out the tree periphery and allow light pene- resulted in no economic benefit (Ferree et al.,
tration to the canopy interior. Even with 1990). Spur pruning produced the smallest
annual follow-up hand-pruning, mechanical increase in fruit size, but also reduced yield
hedging, although successful in maintaining the least, resulting in the greatest cumulative
tree size, has not been economic in maintain- yield and economic value. The outcome of
ing yield and fruit quality (Ferree, 1976b; this study was due in large part to the
Ferree and Lakso, 1979; Ferree and Rhodus, amount and severity of cuts that had to be
1993). In a comparison of pruning six culti- made to achieve the desired canopy shape.
vars annually with limb spreading compared Converting older trees to canopy shapes
with every second or third year without that improve light distribution, such as the
spreading, it was found that the annual treat- palmette leader (Lakso et al., 1989), can be
ment gave the highest absolute return per done with less loss of yield when starting
tree (Ferree and Lakso, 1979). Thus, although with a well-structured central-leader tree
pruning is a costly practice, the cost is justi- (Warrington et al., 1995).
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Pruning and Training Physiology 331

If trees have been neglected and not effects of summer pruning on apple are not
pruned for several years, it is best to make controlled by carbohydrate levels or by the
several large thinning cuts, removing limbs type of carbohydrate present (Taylor and
to open the canopy. It is critical not to prune Ferree, 1986). A redistribution of carbohy-
too excessively in the first year. A few large drates does occur in response to summer
cuts will allow adequate light penetration to pruning and is probably controlled by the
stimulate fruiting wood lower down in the removal of the buds and/or growing points
canopy. In the second and third year, a grad- (Saure, 1987).
ual thinning of the canopy should be done, Quinlan and Preston (1971) showed that
with the focus on renewing the bearing sur- repeated pinching of a growing bourse shoot
face by favouring the retention of young modified the distribution pattern of assimi-
limbs over old ones. If trees have been lates. Movement of photosynthates out of
neglected for more than a couple of years, it the primary leaf towards the growing shoot
is generally not economical to return them to tip was shifted by the pinching to mainly
commercial production. Thus, an economic basipetal transport toward the flower clus-
comparison must be made before deciding ters and the fruitlets, which is the normal
whether to prune or to remove old or direction of transport after terminal bud for-
neglected trees. mation. A similar shift in assimilate distribu-
tion may be responsible for increased fruit
set and improved spur quality in the canopy
14.6 Summer Pruning interior in response to early-summer pruning.

Although summer pruning has been used


for several centuries in some European 14.6.2 Effects on fruits
orchards, the practice was never widely
adapted until more recent times. Saure (1987) Summer pruning has been used primarily to
published a comprehensive review of the improve red fruit colour and fruit quality,
many studies evaluating the impact of the while attempts to control growth by summer
practice on the growth and fruiting of apple. pruning have been less successful. Many
studies show a dramatic improvement in red
fruit colour of difficult-to-colour cultivars,
14.6.1 Effects on physiological processes such as ‘McIntosh’ and ‘Gala’ (Lord and
Greene, 1982; Warrington et al., 1984); how-
Summer pruning affects several of the basic ever, the red colour of cultivars that have a
physiological processes of apple trees (Ferree high colour factor, such as the red strains of
et al., 1984). Photosynthetic rates of basal ‘Delicious’, is often not affected. Summer
leaves remaining after trees were summer- pruning often improves storage quality by
pruned were 11–39% higher than corre- reducing bitter pit, internal breakdown and
sponding leaves on unpruned trees. These water-core (Preston and Perrin, 1974; Myers
rates were equivalent to those in much and Ferree, 1983a). The reduction in these
younger leaves on unpruned trees. As sum- storage disorders is probably related to the
mer-pruning severity increased, the delay in increase in fruit calcium that results from
decline in photosynthesis increased. This summer pruning (Table 14.3). Fruit calcium
delay in the decline of leaf performance was is enhanced by summer pruning early, before
similar to the response induced by increased terminal bud formation and during the time
cytokinin levels. Since the root system of the of active cell division in the fruit, while
summer-pruned trees was not immediately pruning after terminal bud formation has lit-
affected, it might be expected that the root tle effect on fruit calcium concentrations. The
system would continue to produce the same increase in fruit calcium resulting from sum-
quantities of cytokinins as it did prior to mer pruning can often overcome the adverse
pruning, thus increasing the supply to the effect on fruit calcium induced by reduction
remaining leaves. Evidence indicates that the of crop due to frost or excessive growth from
Apples - Chap 14
332

11/4/03
11:01 am
Table 14.3. Effect of summer pruning for 3 years on growth and fruiting of ‘Jonathan’ apple (from Taylor and Ferree, 1984).

Page 332
D.C. Ferree and J.R. Schupp
Marketable fruit (%)
Trunk Canopy
diameter (cm)
area openness1 Firmness Colour Water-
(cm2) (% sky) > 8.0 8.0–7.3 7.3–5.7 (kg) rating2 core3

Control 168a 14b 3.8b 28.8b 65.1a 6.86b 3.8b 0.38b


Summer-pruned 148b 28a 12.7a 48.9a 37.9b 6.98a 4.0a 0.54a
1 Measured by fish-eye lens at tree base.
2 Colour rating: 1 = green to 5 = full red.
3 Water-core rating: 0 = no water-core to 5 = severe water-core throughout cortex.

Values with letters uncommon are statistically different (P = 0.05).


Apples - Chap 14 11/4/03 11:01 am Page 333

Pruning and Training Physiology 333

dormant pruning, but the increase is not ade- vigorous trees that exceed their allotted
quate to overcome a general calcium defi- space. Generally, summer pruning is con-
ciency. If too many shoot leaves are removed ducted on the periphery of the tree remov-
in summer pruning, fruit size can be ing vigorous growth by cutting to the
reduced, as well as fruit soluble-solids con- bud-scale scars or to the first spur on 2-year
centration. Modest summer pruning can wood. Growth produced the year following
achieve the increase in fruit colour without summer pruning has been equal to growth
adverse effects on fruit size and soluble from dormant pruned trees (Myers and
solids; thus, these factors must be closely Ferree, 1983b). Marini and Barden (1982)
monitored to avoid negative effects. found no difference in growth, if vigorous
Although early reports indicated that flow- trees were pruned identically, either while
ering and fruit set were enhanced by summer dormant in winter or while growing in sum-
pruning (Saure, 1987), most recent work mer. Forshey et al. (1992) indicate that mod-
shows either no effect or reductions. Heavy erate summer pruning may have little
summer pruning occasionally causes some practical benefit for vegetative vigour, even
buds to break into growth and flower late in if repeated over several years. They indicate
the season, so that bloom and ripening fruit that summer pruning sufficient to reduce
exist at the same time on trees. This is more vegetative vigour is usually accompanied by
prevalent on some cultivars, such as reduced yield, fruit size and/or fruit
‘McIntosh’ and ‘Jonathan’, and the inflores- soluble-solids concentration and may also
cence is often extended rather than compact. produce significant negative side-effects in
Likewise, the influence of summer pruning flowering and winter-hardiness. Thus, any
on fruit set and fruit abscission is unclear, benefit of summer pruning in controlling
with studies reporting contradictory results. growth is temporary and must be coupled
Since all of these factors influence yield, it is with other techniques and management
not surprising that the reports of the influ- practices to achieve a desirable balance of
ence of summer pruning on yield are variable. growth and fruiting.

14.6.3 Effects on growth 14.6.4 Practical implications

Results on container-grown apple trees indi- In reviewing the research on summer prun-
cate that shoot growth, leaf area, stem diam- ing, several principles evolve that have prac-
eter and root growth are reduced by tical significance (Table 14.4). Pruning early,
summer pruning (Plate 14.2; Taylor and before or just after terminal buds form, can
Ferree, 1981). Summer pruning has been increase current-year fruit set by reducing
advocated as a method of devitalizing very competition from rapidly growing shoots.

Table 14.4. Summary of the influence of summer pruning on the growth and fruiting of apple trees.

Characteristic Increased Decreased No change

Leaf photosynthesis X
Leaf carbohydrates X
Fruit calcium X
Fruit colour X
Bitter pit X?
Water-core X?
Fruit size X?
Fruit soluble solids X?
Late-season flowering X
Growth following season X
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334 D.C. Ferree and J.R. Schupp

Pruning at this time will also result in vege- 20th century, many orchards were planted
tative regrowth in close proximity to the cut at a spacing unsuitable for the rootstock/
and the earlier in the season the pruning is scion/training system combination. As a
done, the greater will be the vigour of the result, overcrowding, decreased production
regrowth. Regrowth in the current season is and lower fruit quality occurred. A means
reduced or eliminated if summer pruning is was needed to control tree growth, and root
performed 1–2 months before harvest but pruning was one of the techniques evalu-
after terminal buds have formed. Regrowth ated. A review of root pruning (Geisler and
in the current season is also reduced as cuts Ferree, 1984b) reported that it was widely
are made into older wood. Cuts made to the practised in European gardens to reduce tree
first spur bearing fruit on 2- or 3-year-old size and promote flowering in the 1800s. In
wood often do the most to open up the later years it was considered too dwarfing to
canopy and promote improved fruit colour, be a recommended practice in either
without undesirable current-season re- European or North American orchards.
growth. Removal of entire shoots by thin-
ning cuts produces little or no regrowth
when done after terminal-bud formation. 14.7.1 Effects on physiological processes
Because the remaining sites of auxin pro-
duction are undisturbed, little disruption of Root pruning of non-bearing apple trees in
the apical-dominance control system occurs. the greenhouse temporarily reduces net
This approach provides the maximum photosynthesis, transpiration and leaf water
improvement in fruit colour since it allows potential (Geisler and Ferree, 1984b; Schupp
more light to reach the fruit. and Ferree, 1990). When roots are removed
Although the primary use of summer by pruning, assimilates are redistributed to
pruning is on fruiting trees to improve fruit the root system, as evidenced by the
colour and quality, there are several other increase in small roots in close proximity to
uses that are beneficial. Removal of water the trunk, coupled with a reduction in leaf
sprouts in the tree centres and on major scaf- and new shoot growth. In a field study, root-
folds of older trees improves light and spray pruned trees had more negative leaf water
penetration and reduces problems from potential, reduced stomatal conductance
some insects. Water-sprout removal can be and reduced transpiration (Schupp et al.,
done any time, but pulling them off early in 1992). There was little effect on concentra-
the season is preferred since cutting later tions of leaf nutrient elements (Schupp and
after the wood hardens leaves basal buds to Ferree, 1987; Schupp et al., 1992).
regrow. Summer pruning can be used on Shoot-tip ethylene production was
young trees to encourage leader develop- reduced at 3, 9 and 17 days after root prun-
ment in such systems as the central leader, ing, while the concentration of the ethylene
slender spindle and central axis (see precursor, 1-aminocyclopropane-1-carboxylic
Chapter 15). Normally it is best to remove acid, in the xylem sap was unaffected. The
several newly developing shoots just below concentration of cytokinin in the xylem sap
the leader as soon as growth is 2–5 cm long. was reduced 1, 3 and 9 days after root prun-
Summer pruning should not be used on ing (Schupp and Ferree, 1994). Thus, ethylene
trees with low vigour. The loss of leaf sur- and cytokinin did not appear to be responsi-
face and the photosynthetic capability it rep- ble for the reductions in growth caused by
resents can only aggravate the pre-existing root pruning.
vigour problem. Although root pruning reduces photosyn-
thesis and transpiration, it has no influence
on the carbohydrate fractions in the leaves or
14.7 Root Pruning shoots, but causes an increase in soluble and
insoluble fractions in the roots (Ferree, 1989).
As orchard intensification became a world- The consistent water-stress effect found from
wide phenomenon in the latter half of the root pruning probably explains most of the
Apples - Chap 14 11/4/03 11:01 am Page 335

Pruning and Training Physiology 335

growth response, as growth rate recovers Compared with young greenhouse-grown


along with regeneration of the root system trees, the effects of root pruning are much
and its ability to meet evapotranspirational greater and of longer duration on bearing
demand. During the interim stress, the tree trees in the orchard. The presence of crop
reduces water use by stomatal closure and and its effect on root regeneration explain
minimizes transpirational surface through the stronger response. For example, the
reduced shoot and leaf growth. The shift in annual increase in trunk cross-sectional area
plant hormones may also play a role directly of fruiting trees was reduced 44% by root
by influencing growth or indirectly by influ- pruning, but only 14% in deblossomed trees
encing water relations. (Schupp et al., 1992). The presence of crop
reduced root regeneration and was, there-
fore, of primary importance for season-long
14.7.2 Effects on growth reduction in vegetative control by root prun-
ing. The resulting long-term water deficit
Results from greenhouse studies on young is thought to be the means by which shoot
apple trees showed that root pruning growth is reduced. Lenz (1986) showed
decreased leaf number per tree, leaf size, that cropping trees increased water usage
total leaf area, shoot growth and decreased compared with non-cropping trees. In a
dry weight of leaves, shoots and roots 6-year root-pruning study, shoot growth of
(Geisler and Ferree, 1984a; Ferree, 1989; ‘Jonathan’ apple was reduced approximately
Schupp and Ferree, 1990). In the field, root 30% each year (Fig. 14.7a), except the single
pruning reduced trunk cross-sectional area, year where rainfall each month of the grow-
shoot length, shoot number to spur ratio, ing season greatly exceeded the long-term
shoot leaf size, and shoot leaf area to spur- average (Ferree, 1992).
leaf area in cropping apple trees (Schupp Root pruning results in root regeneration
and Ferree, 1987, 1988; Ferree, 1992). in close proximity to the cuts. The timing of
The overall growth effects resulting from root pruning affects how much root regener-
root pruning were sufficient to reduce prun- ation takes place in a season. Considerable
ing time and increase within-canopy light root regeneration was evident at the end of
levels. Canopy light penetration and spur the growing season on trees root-pruned
quality were increased. The more severe (the while dormant or at full bloom. Root regen-
closer to the trunk) the root pruning, the eration was less on roots pruned at June
greater were the effects on growth. drop and minimal in preharvest root-pruned

(a) (b)
40 125
a a a
Average shoot length (cm)

35 a a a
30
Yield (kg per tree)

a
100 b
25 a
b b b
20 b
b a a
15 b b
75
10 b
Control b Control
5 Root pruned (bloom) Root pruned (bloom)
0 50
85 86 87 88 89 90 85 86 87 88 89 90
Year Year
Fig. 14.7. Influence of root pruning over 6 years on shoot growth (a) and yield (b) of mature ‘Jonathan’
apple trees. Note the consistent growth effect and reduction in biennial bearing. (From Ferree, 1992.)
Apples - Chap 14 11/4/03 11:01 am Page 336

336 D.C. Ferree and J.R. Schupp

trees (Schupp and Ferree, 1987). Excavations 50 cm, there was no difference in response,
around trees root-pruned annually for 9 as the upper 25 cm of soil has the greater
years showed a reduction in all root sizes. concentration of roots. The optimum time of
Roots < 1 mm in diameter were reduced 20% pruning appears to be around full bloom. In
by root pruning, while the reduction in a study comparing responses of nine root-
larger roots was nearly double this amount stocks or interstock combinations to the
(Ferree, 1994a). The effect of root pruning on effects of root pruning, growth on all trees
root distribution was greatest in the top was reduced 20–30%, with no difference
30 cm of soil, parallel to the location of the whether it was on dwarfing M.9 or vigorous
root-pruning cut at a depth of 25 cm. Roots seedling rootstock (Ferree and Knee, 1997).
below 30 cm were unaffected. Multiple spray applications of low concen-
trations of cytokinins in the field were not
successful in counteracting the reduction in
14.7.3 Effects on fruiting fruit size due to root pruning.
Since root pruning has the potential to
Compared with unpruned trees, root prun- control tree size, a 10-year study evaluated
ing increases yield efficiency (yield per trunk the potential of root-pruning four cultivars
cross-sectional area), fruit colour and soluble- grown in two orchard systems (Ferree and
solids concentration. Preharvest fruit drop Rhodus, 1993). Root pruning was successful
and fruit size are reduced. In large-fruited in containing tree size of apple trees planted
cultivars (‘Melrose’, ‘Golden Delicious’) the at half the recommended in-row spacing.
effect on fruit size was not enough to reduce However, the reduced tree efficiency in the
yields, but in small-fruited cultivars, such as trellis system of dwarf trees or in the central-
‘Jonathan’, yield was reduced (Fig. 14.7b). leader system of semi-dwarf trees resulted in
Root pruning at full bloom increased minimal cumulative yield increase and no
fruit soluble-solids concentration and apparent economic advantage. It was con-
slightly increased fruit firmness but cluded that it is better to plant systems at a
decreased starch hydrolysis at harvest spacing that avoids excessive crowding and
(Schupp et al., 1992). Root pruning does not allows an efficient balance of growth and
appear to affect blossom density or fruit set. fruiting without the need to intervene with
In a long-term study over 9 years, root prun- practices such as root pruning.
ing reduced the biennial bearing pattern. Root pruning is a mechanical means of
Elfving et al. (1991) found that root pruning tree size and vigour control, which can assist
at full bloom in ‘McIntosh’ reduced the inci- in such situations as improper spacing for
dence of stem-cavity browning and brown rootstock or cultivar or loss of crop to frost
core, but not in every year. Root pruning did or following a particularly hard pruning to
not influence ethylene evolution at harvest, bring growth and fruiting balance to trees
but did reduce post-storage ethylene. They (Table 14.5). Under current market condi-
concluded that root pruning at full bloom tions there is a strong emphasis on produc-
was generally less effective than daminozide ing large-sized fruit to satisfy the
in altering fruit behaviour. preferences of apple retailers. Since root
pruning reduces fruit size, it must be
viewed as a ‘rescue treatment’ for orchards
14.7.4 Practical implications with excessive growth and overcrowding,
rather than a standard practice for reducing
Generally, the closer to the trunk that root growth and pruning labour in orchards with
pruning is carried out, the more roots are cut more normal vigour.
and the greater the effect. For bearing trees, It is important to understand the effects of
cuts 70–100 cm from the trunk on two sides root pruning since all trees purchased from a
are required to have the effects on growth nursery have lost a portion of their root sys-
and fruiting previously described (Plate 14.3). tem. Several reports indicate no effect of
In a study comparing depths of 25 and removing most of the roots from nursery
Apples - Chap 14 11/4/03 11:01 am Page 337

Pruning and Training Physiology 337

Table 14.5. Summary of the influence of root pruning on the growth and fruiting of apple trees.

Characteristic Increased Decreased No change

Trunk growth X
Shoot length X
Shoot number X
Spur/shoot ratio X
Shoot-leaf size X
Spur quality X
Pruning time X
Canopy light penetration X
Biennial bearing X
Fruit set X
Fruit yield (no. of fruit) X
Fruit size X
Preharvest drop X
Fruit colour and quality X
Tree-yield efficiency X

trees after they are dug up on subsequent of cells is changed and reaction wood is pro-
tree growth. However, trees propagated duced on the upper side of the branch.
in the field and transplanted under ideal Reaction wood consists of xylem tissue on
conditions caused a 32% reduction in growth one side of the stem that has short, round
compared with trees propagated in the field tracheids and an altered pattern of wall
and not transplanted (Ferree, 1976a). Thus, thickening. Reaction wood can often be
the effect of root pruning can be significant observed on the pruning cut of side limbs,
and care should be taken not to sever roots with the upper half appearing denser than
through cultivation or other cultural prac- the lower half.
tices if no reduction in growth is desired. The stress created by bending has been
shown to result in increased ethylene con-
tent in the internal air spaces within the
14.8 Training branch (Robitaille and Leopold, 1974). The
ethylene reaches a maximum 2 days after
Training utilizes various techniques, includ- bending stress is imposed and returns to
ing pruning, to direct tree growth or form control levels in 3 weeks. A similar increase
and the development of the structural in ethylene was shown to occur by a paste
framework of the tree. Although these forms application of auxin (naphthaleneacetic acid
can be very ornamental (Plate 14.4), gener- (NAA)) to the branch, which in turn caused
ally the tree form fits some predesigned an increase in ethylene. Thus, both auxin
training system described in Chapter 15. and ethylene may play a role in the
Discussion in this chapter will be restricted responses induced by bending.
to the techniques used to control tree growth The growth reduction and increase in
in many orchards. flower initiation caused by bending occur
only during the season when the physical
bending stress is imposed or the limb orien-
14.8.1 Bending tation is changed (Table 14.6). This effect has
commercial value, since, by progressively
Physically bending a branch of an apple tree changing branch orientation by inserting a
results in a reduction in terminal shoot longer spreader each year, a growth reduc-
growth and a redistribution of growth hor- tion can be achieved for several years. The
mones, particularly auxin. The arrangement progressive influence of spreading the stress
Apples - Chap 14 11/4/03 11:01 am Page 338

338 D.C. Ferree and J.R. Schupp

Table 14.6. Influence of bending on apple growth and flowering in the year of treatment (from Ferree,
1994b).

Shoot length Limb area Flowers


Treatment (cm) (cm2) per limb area

Control 26.8a 2.14b 13.2b


Bending 24.4b 2.48a 15.3a

Values with letters uncommon are statistically different (P = 0.05).

over 2–3 years results in the greatest num- The bending of a branch influences not
ber of short, fruitful shoots and the fewest only the amount of growth that occurs
number of undesirable shoots. Cropping in but also the location where shoots arise
successive years has a similar influence on (Fig. 14.8). The influences of auxin begins
the tree, but it is not as predictable as physi- just below the shoot tip and, as the orienta-
cally bending and does not occur until after tion away from vertical increases, the auxin
the limb may have grown for several years influence progresses down the branch and
and stiffened in an upright position, requir- buds on the upper side are released from the
ing considerable force or weight to change hormonally induced apical dominance and
its orientation. The shoot-growth reduction begin to grow. Thus, the more the branch ori-
that results from bending is associated with entation is changed towards the horizontal,
a similar reduction in root growth, so that the greater the number of shoots released
the top-to-root ratio of the young tree is towards the branch base. If basal shoots are
maintained. released due to bending too severely, they
Forshey et al. (1992) suggested that the tend to be very vigorous and upright and
reduced vegetative vigour in limbs that are often require removal during the dormant
bent reduces the production of gibberellins, season. By bending and changing orientation
which are antagonistic to flowering. The for- only 10–15° in any year, shorter shoots are
mation of moderately vigorous lateral shoots produced mostly on the outer half of the
and spurs creates additional sites for flower branch. If excessive spreading or a heavy
formation. These two effects result in crop load on the tip of the branch brings the
increased flowering and earlier fruiting. branch to a horizontal or below-horizontal
Fruit quality is also improved because of position, vegetative growth on the weeping
improved light penetration and because tip portion nearly ceases and several very
more fruit hang free instead of rubbing vigorous suckers will form from buds on the
against the branch on more upright limbs. highest point of the bend. This undesirable

40° 60° 70° 90°

Fig. 14.8. Influence of various degrees of bending on distribution and amount of growth of an apple branch.
Apples - Chap 14 11/4/03 11:01 am Page 339

Pruning and Training Physiology 339

type of response can be avoided by not always found. In training systems such as
bending the limb beyond about 60° from ver- the slender spindle or hybrid tree cone
tical and avoiding overcropping on the tips orchard system (HYTEC), bending is used
of scaffolds of young trees. to weaken the leader and encourage
Several reports indicate that the inclina- growth and fruiting in the lower scaffolds.
tion of the scion, particularly if horizontal, In a study that compared the influence of
reduced growth and increased flowering bending the leader first one way and then
(Tromp, 1972). The study concluded that the the opposite way for different lengths of
reduction of growth and promotion of flow- time, the length of time the leader was bent
ering are independent phenomena and shoot had no effect on shoot growth, but 15 days
orientation interferes in the later phases of appeared to increase flowering, with no
flower formation. The effect on flowering benefit from longer times in the bent posi-
was equivalent to that caused by damino- tion (Ferree and Schmid, 1999). The
zide. A field study over 5 years with ‘Golden hypothesis that the response to bending
Delicious’/M.9 found that inclining the may differ at various times during spring
trunk at planting 45° or 60° from the vertical when growth is initiated was refuted in this
reduced the rate of shoot elongation by 40% study, as no effect was found (Fig. 14.9).
and trunk circumference growth by 17% and Bending in combination with minimal
27%, respectively (Bargioni et al., 1995). Root pruning was successful in reducing growth
total dry weight was reduced by inclination and increasing production of young ‘Fuji’
and roots were more numerous and more trees (Ferree and Schmid, 1994). In a study
elongated in the direction of inclination. The with two cultivars that compared responses
authors propose that the physiological basis on the dwarfing rootstock Mark and the
of the phenomenon may reside in the asym- semi-dwarf rootstock M.7a, bending scaf-
metric distribution of hormones caused by folds increased flowering in the 2 subse-
the inclination. These effects of inclination quent years and increased cumulative yield
probably influence trees trained in such 11% on Mark, but had no effect on the yield
systems as the Lincoln canopy or Tatura of trees on M.7 (Ferree, 1994b). It is unclear
trellis, where trunks or branches are inclined why bending provides an economic benefit
and secured to the trellis framework. by reducing or redirecting growth in some
Practically, bending is accomplished using instances and in others the effects are negli-
many techniques. In central-leader training, gible.
scaffold limbs are spread using wooden,
metal or plastic spreaders, either notched or
pointed to keep them in place. These are nor- 14.8.2 Phloem interruption
mally inserted before growth starts and can
be removed after 4–6 weeks of growth. Others Scoring and ringing, which cause the inter-
prefer to bend by tying limbs down to other ruption of the downward flow of carbohy-
limbs or anchors in the soil or fastening to the drates and hormones in the phloem, have
support system. The ties must be watched been shown to alter growth and fruiting in
and cut if they begin to girdle the limb. apple. The degree or length of time that
Weights have also been used and again care translocation through the phloem is inter-
must be given to avoid girdling. Spreaders rupted is influenced by whether and how
and ties move a limb to a given position, but much of the bark and phloem are removed.
with weights it is more difficult to know Scoring, which is a circumscribing cut
exactly the degree of bending that will occur. through the bark, but not into the wood, has
Special rubber bands can be used to bend the smallest effect. Ringing, which removes
actively growing shoots or a leader, and again different amounts of bark in a circumscribing
the degree of bending is difficult to ascertain. ring, increases in severity with the amount of
Although bending has been commonly bark that is removed until the ultimate of
used to reduce shoot extension and pro- tree death occurs when the tree is unable to
mote flowering, consistent effects were not form new tissue to bridge the ring.
Apples - Chap 14 11/4/03 11:01 am Page 340

340 D.C. Ferree and J.R. Schupp

70

LSD = 11.5
60

Terminal shoot length (cm) 50

40

30
Check
20
Bend
10 Head

Bend + head
0
16 Apr. 25 Apr. 9 May 23 May 6 June 20 June
Time of bending
Fig. 14.9. Interaction between time of bending and heading vigorous laterals on the central leader of ‘Regal
Gala’ on M.7 rootstock (from Ferree and Schmid, 1999).

Scoring and/or ringing interrupt phloem Elfving et al. (1991) found that trunk scoring or
translocation, which causes a build-up of the ringing increased soluble solids and retarded
products of photosynthesis in the portion of loss of flesh firmness before harvest and fol-
the tree above the cut. Growth hormones pro- lowing storage, but had little effect on starch
duced in the apical meristem and young hydrolysis. Scoring or ringing decreased the
leaves are also translocated in the phloem and incidence of brown core, stem-cavity brown-
thus also play a role in the effects produced. ing and scald in some years. Scoring also
The photosynthetic and transpiration rates of reduced post-storage ethylene evolution.
leaves above the scoring cut are reduced for ‘McIntosh’ fruit were longer and had larger
up to 2 weeks, but recover to the control lev- pedicel diameters and weights and more
els of unscored plants in 4 weeks. This effect colour from trees that were ringed (Webster
is thought to be due to the carbohydrate feed- and Crowe, 1969). Results from these studies
back mechanism, which causes photosynthe- were variable on the effects of phloem inter-
sis to slow or stop when carbohydrates build ruption on fruit size and quality and were
up and are not translocated out of the leaves. often not consistent from year to year.
If a ring of bark is removed, the effects on More severe versions of phloem interrup-
photosynthesis last for more than 4 weeks tion (Plate 14.5) can be done with a chainsaw
and a greater reduction in growth would be (Hoying and Robinson, 1992). Chainsaw ring
expected. A number of studies show that girdling involves making two overlapping
phloem interruption reduced terminal horizontal rings, 2 cm deep and spaced 20 cm
growth, trunk cross-sectional area, shoot apart on the trunk. With guillotine girdling,
diameter and number of shoots (Greene and two chainsaw cuts were made to remove all
Lord, 1983; Autio and Greene, 1992). the tissue on opposite sides of the trunk to a
Trunk scoring 19 days after full bloom depth of one-third the radius of the trunk.
reduced fruit size in ‘Cortland’ and ‘Delicious’ Both techniques reduced growth, although not
apples, reduced firmness in ‘Delicious’ and as much as root pruning, with the advantage
increased soluble solids in both cultivars that fruit size was not reduced by girdling
(Greene and Lord, 1978). After storage, (Hoying and Robinson, 1992). Caution must
‘Cortland’ apples from scored trees had a be exercised when applying herbicides under
higher incidence of bitter pit and cork, while trees that have undergone these treatments to
‘Delicious’ fruit exhibited more breakdown. avoid damaging the callus tissue that forms as
Apples - Chap 14 11/4/03 11:01 am Page 341

Pruning and Training Physiology 341

the girdles heal. Furthermore, this treatment the bud, but it was not able to overcome inhi-
has induced the development of heart-rot in bition due to geotropism. Shoots that develop
some trees. It is cautioned that this technique from notched buds usually have a sharper
should be used only to extend the economic crotch angle than shoots that develop from
life of orchards that are nearing the end of buds that were not notched (Verner, 1955).
their useful lifespan (Hoying, 1993). Crotch angle can be improved by using
clothes-pegs or other techniques, as previ-
ously described. Greene and Autio (1994)
14.8.3 Notching found notching consistently effective, indicat-
ing that one could reasonably expect more
Notching is a phloem-interruption technique than 80% of the notched buds to grow into
used to stimulate lateral branching from buds lateral shoots. Notching is a very useful tech-
that would not normally grow. The technique nique for inducing shoots for scaffolds and
removes a 2–3 mm wide strip of bark directly for balancing growth in young trees. In addi-
above a bud. The cut extends down to the tion to using pruning, bending and notching
secondary xylem and around about one-third to induce shoot growth, chemical growth reg-
of the circumference of the stem. Notching ulators (Chapter 17) and the removal of the
stimulates lateral branching by interrupting youngest cupped leaves on the apical meris-
the downward movement of auxin from the tem (Chapter 6) can be used in appropriate
shoot tip (Tamas, 1987). As long as the situations. Enclosing the base of very vigor-
phloem remains severed and auxin is pre- ous upright leaders in plastic sleeves just
vented from reaching the lateral bud, the prior to the beginning of growth has been
growth and development of the bud will be used to stimulate buds to grow (Plate 14.6).
induced. Greene and Autio (1994) found that
the most efficient time to notch was approxi-
mately 2–4 weeks before full bloom. Buds 14.9 Conclusion
notched shortly after bloom grew less than
those notched earlier. Although notching Pruning and training occupy a unique posi-
increased the growth of all bud sizes, the tion in the management of an orchard, being
largest buds were more likely to develop into the most costly cultural practices and the
a lateral shoot and the shoot would be longer practices most often poorly done, with dra-
than from small buds. Notching was most matic influences on orchard efficiency and
effective at inducing shoot growth from buds fruit quality. Pruning interacts with other
on the top of the branch, less effective for cultural practices, such as fertilization, fruit
buds on the side and least effective for buds thinning, growth regulators and pest control.
on the underside of the branch. The percent- Fruit quality is a sensitive indicator of the
age of buds that developed into shoots was effectiveness of pruning, as canopy light dis-
not influenced by wood age or location along tribution influences all aspects of yield and
the length of the shoot. Notching was able to fruit quality. Although good pruning does
overcome bud inhibition imposed by a shoot not guarantee success, economic success
apex by preventing the movement of auxin to without good pruning is inconceivable.

References

Autio, W.R. and Greene, D.W. (1992) Ringing and scoring to control growth of apple trees: results from
1991. Massachusetts Cooperative Extension Service Fruit Notes 57(3), 1–5.
Bargioni, G., Baroni, G., Pietro, T., Pitacco, A. and Ramina, A. (1995) Scion inclination in Malus domestica
Borkh and Prunus spp. influence root growth and distribution. Plant Science 30, 517–520.
Cain, J.C. (1972) Hedgerow orchard design for most efficient interception of solar radiation: effects of tree
size, shape, spacing and row direction. New York Search Agriculture 2(7), 1–14.
Apples - Chap 14 11/4/03 11:01 am Page 342

342 D.C. Ferree and J.R. Schupp

Elfving, D.C. (1990) Growth and productivity of ‘Empire’ apple trees following a single heading-back
pruning treatment. HortScience 25(8), 908–910.
Elfving, D.C. and Forshey, C.G. (1976) Growth and fruiting responses of vigorous apple branches to
pruning and branch orientation treatments. Journal of the American Society for Horticultural Science
101, 290–293.
Elfving, D.C., Lougheed, E.C. and Cline, R.A. (1991) Daminozide, root pruning, trunk scoring and trunk
ringing effects on fruit ripening and storage behavior of ‘McIntosh’ apple. Journal of the American
Society for Horticultural Science 116, 195–200.
Ferree, D.C. (1976a) Effect of rootstocks, propagation method, and transplanting on growth and flower-
ing of young apple trees. Journal of the American Society for Horticultural Science 101, 676–678.
Ferree, D.C. (1976b) Influence of slotting saw mechanical pruning and alar on apple fruit size and quality.
Ohio Agricultural Research and Development Center Research Circular 220, 3–7.
Ferree, D.C. (1978) Influence of heading height of newly planted trees on apple tree growth. Fruit crops –
1978: a summary of research. Ohio Agricultural Research and Development Center Research Circular 239,
5–8.
Ferree, D.C. (1989) Growth and carbohydrate distribution of young apple trees in response to root prun-
ing and tree density. HortScience 24, 62–65.
Ferree, D.C. (1992) Time of root pruning influences vegetative growth, fruit size, biennial bearing and
yield of ‘Jonathan’ apple. Journal of the American Society for Horticultural Science 117, 198–202.
Ferree, D.C. (1994a) Root distribution following 9 years of root pruning ‘Melrose’/M.26 apple trees.
HortScience 29, 76–78.
Ferree, D.C. (1994b) Early performance of two apple cultivars in three training systems. HortScience 29,
1004–1007.
Ferree, D.C. and Forshey, C.G. (1988) Influence of pruning and urea sprays on growth and fruiting of
spur-bound ‘Delicious’ apple trees. Journal of the American Society for Horticultural Science 113,
699–703.
Ferree, D.C. and Knee, M. (1997) No interaction between root pruning and rootstock on growth and per-
formance of ‘Golden Delicious’ apple. HortScience 32, 645–648.
Ferree, D.C. and Lakso, A.N. (1979) Effects of selected dormant pruning techniques in a hedgerow apple
orchard. Journal of the American Society for Horticultural Science 104, 736–739.
Ferree, D.C. and Palmer, J.W. (1982) Effect of spur defoliation and ringing during bloom on fruiting, fruit
mineral level, and net photosynthesis of ‘Golden Delicious’ apple. Journal of the American Society for
Horticultural Science 107, 1182–1186.
Ferree, D.C. and Rhodus, W.T. (1993) Apple tree performance with mechanical hedging or root pruning
in intensive orchards. Journal of the American Society for Horticultural Science 118, 707–713.
Ferree, D.C. and Schmid, J.C. (1994) Influence of training stakes and various pruning and bending tech-
niques on early performance of ‘Fuji’ apple trees. Fruit crops 1994: a summary of research. Ohio
Agricultural Research and Development Center Research Circular 298, 11–17.
Ferree, D.C. and Schmid, J.C. (1999) Influence of temporary bending and heading on branch develop-
ment and flowering of vigorous young apple trees. Fruit crops: a summary of research. Ohio
Agricultural Research and Development Center Research Circular 299, 47–54.
Ferree, D.C., Myers, S.C., Rom, C.R. and Taylor B.H. (1984) Physiological aspects of summer pruning.
Acta Horticulturae 146, 243–252.
Ferree, D.C., Schmid, J.C., Schupp, J.R. and Warrington, I.J. (1990) Influence of pruning treatments on
mature spur-bound ‘Starkrimson Delicious’ apple trees. Fruit crops 1990: a summary of research.
Ohio Agricultural Research and Development Center Research Circular 297, 16–22.
Forshey, C.G., Elfving, D.C. and Stebbins, R.L. (1992) Training and Pruning Apple Trees. American Society
for Horticultural Science, Alexandria, Virginia, 166 pp.
Geisler, D. and Ferree, D.C. (1984a) The influence of root pruning on water relations, net photosynthesis,
and growth of young ‘Golden Delicious’ apple trees. Journal of the American Society for Horticultural
Science 109, 827–832.
Geisler, D. and Ferree, D.C. (1984b) Response of plants to root pruning. Horticultural Reviews 6, 155–188.
Greene, D.W. and Autio, W.R. (1994) Notching technique increases branching of young apple trees.
Journal of the American Society for Horticultural Science 119, 678–682.
Greene, D.W. and Lord, W.J. (1978) Evaluation of scoring, limb spreading, and growth regulators for
increasing flower bud initiation and fruit set of young ‘Delicious’ apple trees. Journal of the American
Society for Horticultural Science 103, 208–210.
Apples - Chap 14 11/4/03 11:01 am Page 343

Pruning and Training Physiology 343

Greene, D.W. and Lord, W.J. (1983) Effects of dormant pruning, summer pruning, scoring and growth
regulators on growth, yield and fruit quality of ‘Delicious’ and ‘Cortland’ apple trees. Journal of the
American Society for Horticultural Science 108, 590–595.
Grochowska, M.J., Karaszewska, A., Jankowska, B., Maksymiuk, J. and Williams, M.W. (1984) Dormant
pruning influences on auxin, gibberellins, and cytokinin levels in apple trees. Journal of the American
Society for Horticultural Science 109, 312–318.
Hoying, S.A. (1993) Benefits and pitfalls of nicking, notching, ringing, girdling and rootpruning apple
trees. Compact Fruit Tree 26, 66–68.
Hoying, S.A. and Robinson, T.L. (1992) Effects of chain saw girdling and root pruning of apple trees. Acta
Horticulturae 322, 167–172.
Lakso, A.N., Robinson, T.L. and Carpenter, S.G. (1989) The palmette leader: a tree design for improved
light distribution. HortScience 24, 271–275.
Lenz, F. (1986) Fruit effects on transpiration and dry matter production in apples. In: Lakso, A. and Lenz,
F. (eds) The Regulation of Photosynthesis in Fruit Trees. Symposium Proceedings, New York State
Agricultural Experiment Station, Geneva, New York, pp. 101–104.
Lespinasse, J.M. (1980) Fruiting habits of apple and how they influence tree forms. Fruit Belge 48(391),
184–198. Translated from French by R.L. Stebbins.
Lord, W.J. and Greene, D.W. (1982) Effects of summer pruning on the quality of ‘McIntosh’ apples.
HortScience 17, 372–373.
Marini, R.P. and Barden, J.A. (1982) Growth and flowering of vigorous apple trees affected by summer
and dormant pruning. Journal of the American Society for Horticultural Science 107, 34–39.
Mika, A. (1975) The mechanism of fruiting inhibition caused by pruning in young apple trees. Fruit
Science Report 2, 32–42.
Myers, S.C. and Ferree, D.C. (1983a) Influence of time of summer pruning and limb orientation on yield,
fruit size, and quality of vigorous ‘Delicious’ apple trees. Journal of the American Society for
Horticultural Science 108, 630–633.
Myers, S.C. and Ferree, D.C. (1983b) Influence of time of summer pruning and limb orientation on
growth and flowering of vigorous ‘Delicious’ apple trees. Journal of the American Society for
Horticultural Science 108, 634–638.
Palmer, J.W. (1989) The effect of row orientation, tree height, time of year and latitude on light intercep-
tion and distribution in model apple hedgerow canopies. Journal of Horticultural Science 64, 137–145.
Preston, A.P. and Perrin, M.A. (1974) The effect of summer pruning and nitrogen on growth, cropping
and storage quality of Cox’s Orange Pippin apple. Journal of Horticultural Science 49, 77–83.
Quinlan, J.D. and Preston, A.P. (1971) The influence of shoot competition on fruit retention and cropping
of apple trees. Journal of Horticultural Science 46, 525–534.
Richards, D. and Rowe, R.N. (1977) Effects of root restriction, root pruning and 6-benzylaminopurine on
the growth of peach seedlings. Annals of Botany 42, 729–740.
Robitaille, H.A. and Leopold, A.C. (1974) Ethylene and the regulation of apple stem growth under stress.
Physiologia Plantarum 32, 301–304.
Rollins, H.A., Jr, Howlett, F.S. and Emmert, F.H. (1962) Factors affecting apple hardiness and methods of
measuring resistance of tissue to low temperature injury. Ohio Agricultural Research and Development
Center Research Circular 901, 71.
Rom, C.R. and Ferree, D.C. (1986) The influence of fruiting and shading of spurs and shoots on spur per-
formance. Journal of the American Society for Horticultural Science 111, 352–356.
Saure, M.C. (1987) Summer pruning effects on apple – a review. Scientia Horticulturae 30, 253–282.
Schupp, J.R. and Ferree, D.C. (1987) Effect of root pruning at different growth stages on growth and fruit-
ing of apple trees. HortScience 22, 387–390.
Schupp, J.R. and Ferree, D.C. (1988) Effect of root pruning at four levels of severity on growth and yield
of ‘Melrose’/M.26 apple trees. Journal of the American Society for Horticultural Science 113, 194–198.
Schupp, J.R. and Ferree, D.C. (1990) Influence of time of root pruning on growth, mineral nutrition, net
photosynthesis and transpiration of young apple trees. Scientia Horticulturae 42, 299–306.
Schupp, J.R. and Ferree, D.C. (1994) Effect of root pruning on shoot tip ethylene production and xylem
concentrations of cytokinin and 1-aminocyclopropane-1-carboxylic acid in young apple trees. Fruit
crops 1994: a summary of research. Ohio Agricultural Research and Development Center Research
Circular 298, 38–42.
Schupp, J.R., Ferree, D.C. and Warrington, I.J. (1992) Interactions of root pruning and deblossoming on
growth, development, and yield of ‘Golden Delicious’ apple. Journal of Horticultural Science 67, 465–480.
Apples - Chap 14 11/4/03 11:01 am Page 344

344 D.C. Ferree and J.R. Schupp

Shigo, A.L. (1990) Tree branch attachment to trunks and branch pruning. HortScience 25, 54–59.
Tamas, I.A. (1987) Hormonal regulation of apical dominance. In: Davie, P.T. (ed.) Plant Hormones and
Their Role in Plant Growth and Development. Martinus Nijhoff, Dordrecht, The Netherlands,
pp. 393–410.
Taylor, B.H. and Ferree, D.C. (1981) The influence of summer pruning on photosynthesis, transpiration,
leaf abscission, and dry weight accumulation of young apple trees. Journal of the American Society for
Horticultural Science 106, 389–393.
Taylor, B.H. and Ferree, D.C. (1984). The influence of summer pruning and cropping on growth and fruit-
ing of apple. Journal of the American Society for Horticultural Science 109, 19–24.
Taylor, B.H. and Ferree, D.C. (1986) The influence of summer pruning and fruit cropping on the carbohy-
drate, nitrogen and nutrient composition of apple trees. Journal of the American Society for
Horticultural Science 111, 342–346.
Tromp, J. (1972) Effects of growth regulation substances and tree orientation on growth and flower bud
formation in apple. Journal of Horticultural Science 47, 525–533.
Tustin, D.S., Hirst, P.M. and Warrington, I.J. (1988) Influence of orientation and position of fruiting later-
als on canopy light penetration, yield and fruit quality of ‘Granny Smith’ apple. Journal of the
American Society for Horticultural Science 113, 693–699.
Tustin, D.S., Cashmore, W.M. and Bensley, R.B. (1998) The influence of orchard row canopy discontinuity
on irradiance and leaf area distribution in apple trees. Journal of Horticultural Science and
Biotechnology 73, 289–297.
Verner, L. (1955) Hormone relations in the growth and training of apple trees. Idaho Agricultural
Experiment Station Research Bulletin 28, 31.
Warrington, I.J., Stanley, C.J., Volz, R. and Morgan, D.C. (1984) Effects of summer pruning on ‘Gala’ apple
quality. Orchardist of New Zealand 57(11), 518–522.
Warrington, I.J., Stanley, C.J., Julian, J.F., Tustin, D.S., Hirst, P.M. and Cashmore, W. (1995) Pruning strate-
gies for restructuring top-dominant central leader ‘Granny Smith’ apple trees. New Zealand Journal of
Crop and Horticultural Science 23, 315–322.
Webster, D.H. and Crowe, A.D. (1969) Effect of gibberellic acid, N-dimethylaminosuccinamic acid (Alar),
ringing and thinning on ‘McIntosh’ apple shape. Journal of the American Society for Horticultural
Science 94, 308–310.

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