Scientia Horticulturae, 30 (1987) 253-282 253

Elsevier Science Publishers B.V., Amsterdam - - Printed in The Netherlands

Summer Pruning Effects in Apple m a Review

MAX C. SAURE
Dorfstr. 17, D-2151 Moisburg ( F.R.G.)
(Accepted for publication 16 October 1986)

ABSTRACT

Saute, M.C., 1987. Summer pruning effects in apple - - a review. Scientia Hortic., 30: 253-282.

Summer pruning effects in apple have generally been attributed to improved light penetration
and to reduced carbohydrate supply. However, the common basis to most of these effects seems
to be the retardation of senescence, a process under hormonal control. Summer pruning causes a
temporary loss of apical dominance except when thinning cuts are used. It also causes a temporary
increase in cytokinin supply, presumably mainly by increased export from the roots. Both effects
presumably result from reduced auxin availability.
Depending on its timing, the resulting rejuvenation may consist of mobilization and redistri-
bution of nutrients and phytohormones, breaking of axillary buds, inhibition of flower induction,
delayed fruit development and later induction of dormancy. The extent of pruning responses
increases with the vigour of the tree, the earliness of pruning and its severity. Heading and stub-
bing are more effective in provoking these responses than thinning of shoots. In areas with a short
season, summer pruning may not only delay, but also prevent, the onset of dormancy, with adverse
effects on winter hardiness. The mostly positive influence of summer pruning on fruit colour by
factors in addition to better light penetration is discussed. Summer pruning, where considered
necessary, needs to be adjusted to local conditions and coordinated with other cultural practices.

CONTENTS

1. Introduction
2. Results of summer pruning
2.1. Modification of growth
2.2. Modification of yield
2.3. Modification of flowering
2.4. Modification of fruit development
3. Sources of variation in summer pruning effects
3.1. Selection of standards
3.2. Pruning procedures
3.3. Environment
3.4. Tree characteristics

0304-4238/87/$03.50 1987 Elsevier Science Publishers B.V.

254

4. The physiology of plant responses

4.1. Removal of leaves
4.2. Removal of buds and shoot tips
5. Summer pruning as an integrated cultural practice
Acknowledgements
References

1. INTRODUCTION

Summer pruning has been known to European apple growers as a cultural

practice since about the middle of the 17th century (Kemmer, 1948). However,
it was widely neglected for several decades of the 20th century, when commer-
cial apple production expanded rapidly and fruit science became established.
Most probably, this is due to the partially inconsistent effects of summer prun-
ing, as described by Gardner et al. (1952) in their excellent review of the earlier
literature.
Recently, summer pruning has regained the attention of apple growers, as a
means of:
- -limiting the size of trees, especially in modern high-density plantings;
- - improving fruit development in these for better fruit finish and better stor-
age quality.
Recent experimental work has been summarized by Kluge (1983), Barden
and Marini (1984) and Ferree et al. (1984). These investigations have not
unequivocally confirmed the benefits of summer pruning in overcoming the
negative side-effects of modern highly intensive production methods. The rea-
sons for the diverse and partially contradictory results are still not fully under-
stood. Without such an understanding, it remains very difficult to decide
whether summer pruning should be done as an integrated part of standard
cultivation methods, and if so, how to adapt it properly to different growing
conditions with respect to timing, method and .degree of pruning.
This review tries to reconcile the contradictions between results of different
investigations and to identify a common basis of various pruning responses. It
is hoped that suggestions resulting from it will encourage further research. The
nuclear physicist W. Heisenberg (1973) pointed out that one of the benefits
of new theoretical approaches is that they may permit the finding of new facts,
and known facts may appear in a new light. A new approach to summer pruning
is needed.
This review is confined to recent research results. For earlier publications,
it refers to the extensive work of Gardner et al. (1952), comprising 95 refer-
ences. The term "summer pruning" will be used in its widest sense, i.e. the
removal of leafy branches, shoots or parts thereof, irrespective of the timing,
method or severity. The different types of pruning procedures will be dealt with
in Section 3.2.
 255

2. RESULTS OF SUMMERPRUNING

2.1. Modification o/growth

Gardner et al. (1952) stated that summer pruning does not necessarily have
either a dwarfing or an invigorating influence, and therefore does not generally
retard growth more than dormant pruning. However, during the last decade,
summer pruning has been recommended repeatedly as a proper practice to
reduce tree vigour, e.g. in Italy (Innerhofer, 1974), The Netherlands (Lem-
mens, 1975, 1977), Germany (Utermark, 1976) and the U.S.A. (Stembridge,
1979).
Indeed, in most experiments summer pruning reduced stem thickening,
although there was great variability in the extent of reduction (Lord et al.,
1979a; Marini and Barden, 1982a, e; Nelgen, 1982; S~ik5 and Laurinen, 1982;
Greene and Lord, 1983; Mika et al., 1983; Myers and Ferree, 1983b; Polefska,
1983; Ferree et al., 1984; Domoto, 1985; Saure, 1985a; Wagenmakers, 1985).
Subsequent shoot growth (regrowth) in the season of pruning was often lim-
ited, and the canopy was thus drastically reduced. Leaves of the regrowth were
generally small (Maggs, 1965; Taylor and Ferree, 1981; Ferree et al., 1984a)
and contained less chlorophyll a and b ( Singha and Baugher, 1985 ). The sus-
ceptibility of the wood to winter injury sometimes increased substantially com-
pared with unpruned or dormant-pruned trees (Lord et al., 1979a; Domoto,
1985; Saure, 1985a). All these effects have been interpreted as devitalization.
However, these results do not confirm a general dwarfing effect. As an imme-
diate response to summer pruning, growth rate of the remaining shoots may
be enhanced, and termination of growth is often delayed. Leaves left on the
tree turn dark green, and their abscission is postponed ( Maggs, 1965; Lord et
al., 1979a; Taylor and Ferree, 1981; Marini and Barden, 1982b; Nelgen, 1982;
S~ik5 and Laurinen, 1982; Myers and Ferree, 1983a; Polefska, 1983). In the
year after summer pruning, shoot growth is generally not restricted or is even
more vigorous (Lord et al., 1979a; Prigge, 1981; Marini and Barden, 1982a, e;
Nelgen, 1982; Greene and Lord, 1983; Mika et al., 1983; Myers and Ferree,
1983a; Taylor and Ferree, 1984a; Wagenmakers, 1985). Saure {1985a) noticed
a reduced total growth increment of young apple trees in the season after severe
pruning, but length of individual shoots was increased. The increased shoot
length was partially due to a larger number of internodes, but even more to
longer internodes. Most of these observations point to a redistribution of growth
rather than to devitalization, and Myers and Ferree (1984) interpreted sum-
mer pruning as a localized, invigorating influence.
Summer pruning often results in hastened bud break of axillary buds,
depending on pruning method and plant vigour (cf. Section 3). Lord et al.
(1979a) observed this proliferation of growing points mainly below the cuts
on terminal and vigorous upright shoots. Marini and Barden (1982a), Saure
256

(1985a) and Wagenmakers (1985) reported that bud break on apple trees
pruned in August or September, respectively, started earlier the next season
as compared to dormant-pruned trees. Flower bud opening in spring may be
also hastened (Preston and Petting, 1974; Marini and Barden, 1982a; Taylor,
1982; Link, 1984; Taylor and Ferree, 1984a). This could explain why Engel
and Lenz (1981) found flower buds in summer-pruned trees to be more sus-
ceptible to late spring frosts. However, Morgan et al. (1984) did not find this
influence on flower bud opening in spring, and Link (1984) reported that sum-
mer pruning increased frost survival of flower buds. Occasionally, summer
pruning may induce flowering in autumn (Lord et al., 1979b ). In regions with
warm winters, bud break may be delayed by summer pruning, due to an
increased chilling requirement of the buds (Chandler and Brown, 1957).
In the season after treatment, the previous promotion of lateral bud devel-
opment by early summer pruning may cause a more favourable branch orien-
tation in certain cultivars (Aselage and Carlson, 1977). However, Myers and
Ferree (1986) did not succeed in distributing vigour more evenly along the
limb and away from the distal section. According to Gardner et al. (1952), light
summer pruning tends to encourage fruit spur development.This has also been
observed by Lemmens (1977), Stembridge (1979), Belter and Thomas (1980)
and Domoto (1985). Myers and Ferree (1983c) noted that summer pruning,
under certain conditions, may prevent the decline in the amount of leaf area
in spurs which usually occurs in the interior of unpruned trees. Such effects
are not limited to apple but have also occurred in sweet cherries (Prunus
avium), with more fruiting wood in the lower part of the canopy (Widmer and
Zbinden, 1984), and in peach (Prunus persica), with more but less vigorous
growing points (Rom and Ferree, 1985). Stiles (1980) agreed that summer
pruning stimulates lateral shoot growth, but reported flower formation was
reduced (cf. Section 2.3).

2.2. Modification of yield

One of the objectives of summer pruning is to get higher, earlier and more
regular yields,as expressed in the proverb cited by Gardner et al. (1952) "Prune
in winter for wood and in summer for fruit".Link (1984) noted that trees that
were summer-pruned rather than dormant-pruned always produced highest
yields. In the long run, summer-pruned trees yielded about 25% more than
dormant-pruned trees. However, he concluded that the beneficial effects of
pruning exclusively in the summer are not the immediate result of summer
pruning, but of avoiding pruning in the previous winter. Lemmens (1979, 1982)
consistently reported higher yields after summer pruning, especially in the tri-
ploid 'Karmijn de Sonnaville'. Quinlan and Preston (1971) noticed that reg-
ular pinching of bourse-shoots, starting shortly after full bloom, resulted in
increased yields. Pinching may further cause earlier yields by reducing exces-
 257

sive shoot growth and favouring spur formation, as observed by Zbinden and
Widmer (1980) in cherry and plum.
Other authors could not confirm a consistent effect of summer pruning on
cropping (Preston and Perring, 1974; Marini and Barden, 1982d; Jackson et
al.,1983; Myers and Ferree, 1983b; Ferree, 1984; Bootsma, 1984; Domoto, 1985;
Wagenmakers, 1985), and Belter and Thomas (1980) were unable to make
apple trees more precocious. In many of the more recent experiments, summer
pruning even reduced yield under certain conditions (Bos, 1974; Engel, 1974;
Borsboom, 1976; Aselage and Carlson, 1977; Belter and Thomas, 1980; Brun-
ner and Droba, 1980; Stiles,1980; Prigge, 1981; S~ik5 and Laurinen, 1982; Sus
and Prskavec, 1983; Akkerman, 1984; Taylor and Ferree, 1984a; Williams, 1985)
and depending on the standards selected. Taylor (1982) got a great reduction
of yield on a per-tree basis,but not per unit of canopy volume.
Since "yield" is a very complex subject, the obvious contradictions men-
tioned above need to be analyzed in order to get a more general understanding
of these summer pruning effects.In principle, low yields may be the result of
reduced flowering or of impaired fruitdevelopment.

2.3. Modification of flowering

Occasionally, flowering in apple may occur from lateral buds of the last sea-
son's extension growth. More regularly, flowering occurs from terminal buds
on spurs and medium-long shoots. In any case, formation of flower buds
requires:
- induction of flower initials;
-

- proper development of these flower initials.

-

However, very few reports dealing with the influence of summer pruning on
subsequent flowering reveal the way in which flowering may have been affected.
Gardner et al. (1952) concluded from the literature that summer pruning
may promote flower initiation if done early enough and with the appropriate
pruning method. Later investigations have confirmed a beneficial effect of
pinching on flower bud formation (Mika et al., 1983 ). Lord et al. (1979a) also
reported that the ability of secondary shoots to form flowers was increased the
earlier pruning was done. Several authors reported an increased formation of
fruit spurs after summer pruning (cf. Section 2.1). Taylor (1982) noticed a
higher proportion of inflorescences on terminal spurs, and Link (1984) found
a higher flowering density, probably resulting from the larger number of spurs.
Conversely, Myers and Ferree (1983c) reported fewer fruit spurs but more
flowers per cluster the year after summer pruning, indicating an invigoration
of flower buds. Marini and Barden (1982a) noted a swelling of the terminal
buds below the pruning cuts, but most of these buds, although resembling flower
buds, did not bloom. A promotion of flowering following summer pruning has
258

been claimed, especially in less vigorous trees (Dietz, 1984 ) and on more basal
parts of shoots (Lemmens, 1977).
Taylor and Ferree (1984a) could not generally confirm an influence of sum-
mer pruning on flower cluster density and quality, and Lord et al. (1979a) did
not recommend summer pruning for this purpose because of unpredictable
results. Greene and Lord (1983), Morgan et al. (1984) and Wagenmakers
(1985) did not find consistent effects of summer pruning on flowering. Reduced
flowering, under certain conditions, has been reported by Ferree and Stang
(1980), Nelgen (1980), Stiles (1980), Engel and Lenz (1981), Lemmens
(1982), Marini and Barden (1982a) and Mika et al. (1983).

2.4. M o d i f i c a t i o n o / f r u i t d e v e l o p m e n t

A modification of fruit development by summer pruning may influence both

yield and quality. Belter and Thomas (1980) did not find a single fruit after
harvest from flower buds differentiated adjacent to summer pruning cuts. They
assumed that in the distal buds, the flower initials may have been incompletely
developed. Also, Marini and Barden (1982a) noted in 'Delicious' that none of
the few distal buds induced to flower as a result of summer pruning ever pro-
duced a fruit. Ogata et al. (1986) reported that after summer pruning, some
flowers developed terminally on secondary shoots and produced abnormally-
shaped fruits.

F r u i t set. ~ Fruit set was not influenced, or in some instances it was reduced,
in experiments by Lord et al. (1979a). It was definitely diminished in a study
by Link (1984), and there was a tendency towards reduced fruit set after early
summer pruning done by Myers and Ferree (1983c). Pruning treatments did
not affect fruit set in experiments by Greene and Lord (1983) and by Taylor
and Ferree (1984a, b ). Both shoot removal and shoot tip removal starting 15
days after full bloom increased initial fruit set (Quinlan and Preston, 1971 ).
Furthermore, Ferree and Stang (1980) stated that fruit set can be increased
by summer pruning during the first half of the growing season, but that there
was no effect the year after pruning.

Reports on the influence of summer pruning on fruit abs-

F r u i t abscission. ~
cission are as contradictory as they are on fruit set. However, it should be noted
here that a clear distinction between the two is not always possible, as some
authors have reported only the percentage of fruit set at harvest. This does not
explain whether only the initial fruit set, or also the fruit drop at certain periods
thereafter, was involved. Quinlan and Preston (1971) observed that early
removal of complete bourse-shoots resulted in fewer fruits at harvest, although
it promoted fruit set, whereas repeated pinching of the bourse-shoots increased
the number of fruits retained to harvest. The reduction of fruit abscission by
 259

early pinching has been confirmed by Bos (1974), and also by Varga (1971)
in pears. Promotion of June drop by severe pruning was reported by Lord et
al. (1979b). Link (1984) found fewer fruits per cluster on summer-pruned
trees, with a large percentage of clusters shedding all their flowers or fruitlets.
Summer pruning consistently reduced pre-harvest drop in 'Delicious' and
'Stayman', probably as a result of delayed ripening (Marini and Barden, 1982d;
Barden and Marini, 1984). Previous season's summer pruning reduced the
effect of NAA thinning in years with a generally good fruit set, probably due
to an increased proportion of terminal flower buds (Taylor and Ferree, 1984b).

Fruit s i z e . ~ Fruit size is often seriously reduced after summer pruning (Per-
ring and Preston, 1974; Terblanche et al., 1977; Lord et al., 1979b; Van der
Boon, 1980; Brunner and Droba, 1980; Janse, 1980; Myers, 1981; Marini and
Barden, 1982d; Nelgen, 1982; Ferree, 1984; Link, 1984; Domoto, 1985; Wil-
liams, 1985). However, this effect may vary depending on the time, method
and severity of pruning, on the year, and on the cultivar (cf. Section 3 ). Some
authors found no tendency towards smaller fruits (Schumacher et al., 1974;
Lord and Greene, 1982; Miller, 1982; Wagenmakers, 1985). There are only few
reports on increased fruit size (Engel, 1974; Silk5 and Laurinen, 1982; Taylor
and Ferree, 1984b, 1986), and this was probably the result of a reduction in
yield caused by summer pruning rather than a direct influence. Little infor-
mation is available as to the causes of fruit size reduction. Nelgen (1982) stated
that summer pruning did not visibly influence cell number, but late summer
pruning slightly reduced cell size.

Maturation. ~ Delayed development of the fruits could also affect maturation.

Marini and Barden (1982d) noted a delayed conversion of starch, especially
in fruit from the interior of the canopy. They considered the delay in fruit
starch depletion ~ like suppressed pre-harvest drop - - to be an indicator of
delayed maturation. This effect is not limited to apple. Protracted maturation
has also been observed in peach after topping the trees 3 weeks prior to harvest
(Johnson and Coston, 1985). Struklec (1981) noted less chlorophyll reduction
in apples during the whole storage season, and Nelgen (1982) observed that
the skin and flesh of apples from summer-pruned trees were greener. However,
fruit background colour, considered to be a reliable maturity index for apples,
was not significantly affected in pruning experiments by Morgan et al. (1984).
On the contrary, higher rates of ethylene evolution of maturing fruits on sum-
mer-pruned trees have provided some evidence that summer pruning hastened
maturity of the apple fruit, but other maturity factors were not affected in
these apples (Taylor and Ferree, 1984b).

Soluble s o l i d s , m Marini and Barden (1982d) observed that increase of soluble

solids was prevented within 2 weeks after summer pruning. A lower content of
260

soluble solids at harvest time has been confirmed by many other authors, e.g.
Perring and Preston (1974), Lord et al. (1979b), Lemmens (1980), Nelgen
(1980, 1982), Myers (1981), Prigge (1981), Barden and Marini (1984) and
Taylor and Ferree (1984a). Again, like other responses to summer pruning,
the reduction in soluble solids may be inconsistent, for several reasons (cf.
Section 3).

~ No consistent effect of summer pruning on acidity has been observed.

Acidity.
While acidity was generally increased in experiments by Struklec (1981), Nel-
gen (1982) got a partial decrease, especially if pruning was done in late sum-
mer, and no significant difference was found by Perring and Preston (1974).

Firmness. ~ Fruit firmness, another factor showing some relation to matura-

tion and ripening, also does not respond unequivocallyto summer pruning. No
effect was observed by Miller (1982), Marini and Barden (1982d), Greene and
Lord (1983) and Taylor and Ferree (1984a), whereas Myers and Ferree (1983b)
and Nelgen (1980, 1982) sometimes found reduced firmness, but only after
pruning late in summer. Increased firmness of fruit flesh in apple was noted
by Lemmens (1975), Lord et al. (1979b) and Struklec (1981), and in peach
by Johnson and Coston {1985).

Red c o l o u r . ~ Gardner et al. (1952) reported that the positive influence of

summer pruning on the development of red colour in the skin of apples and
other fruits depended on the amount of sunlight reaching the fruit directly.
Since most fruits develop their colour shortly before ripening, they concluded
that pruning may be done relatively late in the season to enhance colour. A
promotion of red colour by summer pruning has since been confirmed by many
authors, such as Bos (1974), Engel (1974), Preston and Perring (1974), Lord
et al. (1979b), Belter and Thomas (1980), Lemmens and Spruit (1980), Janse
(1980), Bootsma (1984), Morgan et al. (1984) and Williams (1985). How-
ever, the effect may vary, depending on the year (Miller, 1982; Taylor and
Ferree, 1984a) and on the treatment (Lord and Greene, 1982). Sometimes,
summer pruning had no effect on colour formation (Greene and Lord, 1983;
Myers and Ferree, 1983b; Wagenmakers, 1985 ), and sometimes its effect was
even negative after early pruning of a difficult-to-colour cultivar like 'Jona-
gold' (Gebert et al., 1986). Since blush and maturity are not closely related,
an increase in red colour does not necessarily indicate an acceleration of
maturation (Morgan et al., 1984).

Skin f i n i s h . ~ Skin finish contributes to the attractiveness of apples. Lem-

mens (1979) pointed out that summer pruning may reduce russeting in the
triploid 'Karmijn de Sonnaville', which often suffers from severe russeting
around the calyx. However, he later reported that pruning too early may increase
 261

russeting (Lemmens, 1982). This also holds true for very severe summer prun-
ing (Bos, 1974 ). Russeting was not influenced by summer pruning treatments
in 'Golden Delicious' and 'Melrose' (Ferree, 1984).

Storage quality. ~ Summer pruning is often considered to have a beneficial

effect on storage quality. This has been confirmed, to varying degrees, for:
bitter pit (Bangerth and Link, 1972; Preston and Perring, 1974; Lemmens,
1975, 1982; Borsboom, 1976; Terblanche et al., 1977; Lord et al., 1979b; Van
der Boon, 1980; Nelgen, 1980; Schumacher et al., 1980; Struklec, 1981; Mar-
ini and Barden, 1982b; Myers and Ferree, 1983b; Link, 1984; Kluge, 1985);
- - internal breakdown ( L e m m e n s , 1975,1982; Lord et al., 1979b; Van der Boon,
1980; Struklec, 1981; Nelgen, 1982);
m w a t e r core (Myers, 1981; Marini and Barden, 1982b; Myers and Ferree,
1983b);
--rotting:lenticel rot, gloeosporium, and unspecified (Preston and Perring, 1974;
Lemmens, 1975, 1982; Nelgen, 1982).
However, bitter pit was not reduced in experiments by Utermark (1976),
Greene and Lord (1983) and Sus and Prskavec (1983), or if summer pruning
was only light (Terblanche et al., 1980) or rather late (Myers and Ferree,
1983b). Sometimes, summer pruning reduced bitter pit only in combination
with calcium sprays (Schumacher and Fankhauser, 1967; Engel, 1974), and
occasionally it even increased bitter pit, especially if done early or heavily
(Schumacher and Fankhauser, 1972; Schumacher et al., 1974). Internal
breakdown was not affected in experiments by Schumacher et ah (1974), Uter-
mark (1976), Lord and Greene (1982) and Greene and Lord (1983). A ten-
dency towards increased incidence of water core after summer pruning instead
of a reduction was noted by Taylor and Ferree (1984a).
A lower incidence of bitter pit can often be related to a reduction in fruit
size, which may result from summer pruning (Schumacher et al., 1980; Ter-
blanche et al., 1980). Moreover, the occurrence of this and other physiological
disorders has also been attributed to reduced levels of Ca, or to increased levels
of K and Mg in the fruits, i.e. to an increased (K+Mg)/Ca ratio. This view
finds some support by observations that summer pruning
- raised the level of Ca in apple fruits (Perring and Preston, 1974; Terblanche
-

et al., 1977; Nelgen, 1980; Struklec, 1981 (but not in the first year); Lem-
mens, 1982; Link, 1984), or
--lowered the K or K + M g level in the fruits (Perring and Preston, 1974;
Lemmens, 1982; Nelgen, 1982; Link, 1984).
However, Borsboom (1976), Terblanche et al. (1980), Lord and Greene
(1982), Marini and Barden (1982d), Nelgen (1982), Greene and Lord (1983),
Kluge (1985) and Taylor and Ferree (1986) found no, or inconsistent, effects
of summer pruning on fruit Ca. Perring (1985) observed that late summer
pruning increased the redistribution of Ca from the core to outer fruit parts
262

during storage. Taylor and Ferree (1986) reported that summer pruning
increased K concentration in harvested fruit, but Struklec {1981 ) concluded
from her experiments that differences in fruit K may be attributed to differ-
ences in yield rather than to direct effects of summer pruning.
In the leaves, increased Ca levels as a result of summer pruning have been
recorded by Utermark (1976), Struklec (1981) and Jang and Ko (1985), in
the lower leaves in particular by Van der Boon (1980), and in spur leaves in
general by Taylor and Ferree (1986). However, in leaves of shoot regrowth
after summer pruning, the level of Ca and Mg was lowered, but the level of K
and N was increased (Taylor and Ferree, 1986 ). Reduced levels of leaf K have
been noted by Utermark (1976), and Nelgen (1982) observed that leaf Ca may
be reduced after early summer pruning but leaf K was not affected.

3.S O U R C E S OF VARIATION IN S U M M E R P R U N I N G EFFECTS

There is a great diversity of tree responses to summer pruning, with many

contradictions still not resolved. For a better understanding, an identification
of the underlying mechanisms is required. Lord et al. (1979a) emphasized that
the results of many pruning experiments are difficult to interpret for lack of
sufficient experimental description or statistical design. Marini and Barden
(1982a) also complained that evaluation and comparison of the early studies
are difficult because of different locations, poor design or lack of statistical
analysis, different treatments, and incomplete descriptions.

3.1. Selection of standards

One of the most important causes of differences in the results and conclu-
sions of pruning experiments is the use of inappropriate standards. Most recent
investigations used standards where either summer pruning differed in method
and/or severity of dormant pruning, or the control trees were not pruned at
all, or summer pruning was compared to dormant summer pruning, thus
causing differences in the amount and type of wood removed by the respective
treatments. This appears feasible where only physiological processes triggered
by summer pruning are to be investigated, or where the only objective of prun-
ing is to improve fruit quality immediately by better exposure to light. How-
ever, where the influence of pruning season on the whole plant system, i.e. on
growth responses, flowering and fruiting, is to be investigated, Saute (1980)
postulated that no other factor except timing should be altered, as one equation
with two or more unknowns cannot be solved. Accordingly, Myers and Ferree
(1984) pointed out that the interpretation of summer pruning effects on tree
size control can be strongly influenced by the control that is used as basis for
comparison, and by the point of time that is selected for evaluation.
Actually, only few studies compare the effect of season without altering other
 263

variables (e.g. Prigge, 1981; Marini and Barden, 1982a,b,d,e; Mika et al., 1983;
Saute, 1985a), but even if identical dormant pruning was selected as standard,
the results may differ depending on the basis selected for comparison. By sum-
mer pruning, Taylor (1982) got a yield reduction per tree, or per ha, but not
per m 3 canopy volume (cf. Section 2.2). Another important factor is whether
dormant pruning in the previous winter or that in the next winter serves as
control. The appropriate standard will depend mainly on the timing of summer
pruning and on the general objectives of the experiment.
The proper timing of evaluation is also essential to separate direct and indi-
rect pruning effects. In many experiments, they were not distinguished prop-
erly. This problem is aggravated when experiments are continued for more
than one season. Saure (1981, 1985a) has pointed out that pruning predomi-
nantly causes short-term effects which are compensated by feed-back reac-
tions of the plant, and may additionally be disturbed by the cultural practices
they provoke. For example, it has been shown that light interception may be
improved in the year of summer pruning, but may be reduced in the next year
because of vigorous regrowth (Porpiglia and Barden, 1981) - - which fruit
growers tend to remove early. Similarly, fruit size, which is generally reduced
by summer pruning, may be larger in the following season because of reduced
yield (cf. Section 2.4). Because of the wave-like course of plant responses
(contrary to the often expected linear responses), pruning experiments need
permanent, concrete observations and records. Instead of being extended
beyond the following season, they ought to be repeated with new, uniform
material in order to assess the effects of uncontrollable factors such as light,
temperature, rainfall and crop load.

3.2. Pruning procedures

Summer pruning may be done in many different ways with respect to timing,
method and severity.

Timing. m Gardner et al. (1952) concluded that summer pruning may have
quite different effects depending on its timing. They suggested that very early
summer pruning was most effective in "diverting the energy of the plant into
other developing or already developed tissues", resulting in " . . . a more effi-
cient functioning of the remaining tissues". It was also most effective in
encouraging fruit-spur development. Pruning later during the period of most
rapid vegetative growth resulted in one of the following effects:
it may lead to a greater accumulation of nutrients, with beneficial effects on
the formation of flower buds, especially on spurs and on the basal and median
portions of the growing shoots;
264

- - i t may result in the opposite effect, through the consumption of nutrients,

by forcing axillary buds into growth, leading to the formation of secondary
shoots instead of flower buds.
They also suggested that pruning late in the season nearly always promoted
the accumulation of nutrients in the remaining shoots, since no new growth
would take place to utilize the stored foods. However, this accumulation would
occur too late to benefit flower bud initiation. Regarding fruit development,
Gardner et al. (1952) mentioned that pruning could be done rather late not
only for better fruit colour, but also for less sunburn of the fruit, which is likely
to occur after early summer pruning.
Some of the more recent publications further investigate the importance of
timing. Myers and Ferree (1983c) observed that in late-pruned trees the point
at which lateral shoot growth begins on vertical shoots of the previous season
is lower than in early-pruned trees. There are some reports that early summer
pruning may be beneficial for spur development and subsequent flower bud
formation in the same year (Lord et al., 1979a; Domoto, 1985; Ogata et al.,
1986), but other reports show that flowering is rather negatively affected when
summer pruning is done early (Nelgen, 1980; Lemmens, 1982; Dietz, 1984;
Domoto, 1985). However, early summer pruning may improve fruit set (Quin-
lan and Preston, 1971; Ferree and Stang, 1980). In certain cases, the incidence
of bitter pit and internal breakdown was reduced when summer pruning was
done early (Schumacher et al., 1974 ), but russeting was increased (Lemmens,
1982). Some authors found more adverse effects of early summer pruning on
fruit size as compared to later pruning (Lemmens, 1982; Myers and Ferree,
1983b; Morgan et al., 1984), but fruit growth was more reduced by late than
by early summer pruning according to observations by Domoto (1985) in apple,
which agrees with findings by Akkerman (1984) in pear. Schumacher et al.
(1974) found no specific effect of timing within growing season on fruit size.
The reduction of soluble solids is generally greater after late than after early
summer pruning (Nelgen, 1980; Morgan et al., 1984; Ogata et ah, 1986),
although according to Gardner et ah (1952), the timing of pruning generally
had little influence on maturation.

Method. ~ In principle, summer pruning may consist of the following methods:

(1) removal of the apical parts of shoots, either
green tips ("pinching"), or
- - a portion of more lignified shoots ("heading"), or
upper parts of branches, leaving stubs of older wood ("stubbing");
(2) removal of complete shoots or branches back to their point of origin, or
back to an apically dominant side branch/side shoot ("thinning").
This can be done at the periphery or in the interior of the canopy, and the
various methods may be used separately or in combination.
It appears from the earlier literature (Gardner et al., 1952) that summer
 265

heading tends to stimulate vegetative growth, thus reducing flower bud for-
mation, whereasthinning, under certain conditions, may encourage fruit spur
formation and promote flower bud formation. Pinching has the reputation of
promoting spur and flower formation, but frequently has not provided the
expected response unless it is followedby successive pinching of secondary and
tertiary shoots. More recently, Lord et al. (1979a) noted that the effects of
pinching and heading on regrowth were comparable in some instances, but
heading frequently caused more regrowth, fewer flowers and less stem thick-
ening than did pinching. On the other hand, Miller (1982) considered stubbing
to be more devitalizing than heading because stubbing resulted in significantly
less regrowth of either current season's or last year's shoots. Comparing head-
ing and thinning, Mika et al. (1983) recorded no difference in the number of
flower buds or yield of fruit when the same amount of wood was removed.
Prigge (1981) found heading of all shoots in summer to be more detrimental
to fruit size than dormant-type summer pruning consisting mainly of branch
thinning. June-drop was reduced by repeated pinching of bourse-shoots, but
was increased by complete removal of bourse-shoots (Quinlan and Preston,
1971).

S e v e r i t y . ~ Gardner et al. (1952) reported that increasing severity of summer

pruning increasingly induced breaking of axillary buds, thus theoretically
causing a dissipation of stored nutrients. This would cause a weakening of the
remaining shoots (and perhaps the whole plant) as they entered the dormant
season less richly supplied with stored nutrients. This would certainly not pro-
mote the formation of fruit spurs or flower buds, as was sometimes observed
after light early shoot thinning. Rom and Ferree (1985) increasingly reduced
leaf, shoot, root and total dry weight by increasing pruning severity in young
peach trees which have some responses to summer pruning treatments in com-
mon with apples (Rom and Ferree, 1984). Saure (1985a) pointed out that
young apple trees pruned very severely suffered most from early winter frosts.
The physiologicallystimulating effect of severe summer pruning has been con-
firmed by Ferree et al. (1984), who observed higher transpiration and photo-
synthesis rates with increasing severity of summer pruning.
With respect to fruit development, Bos (1974) warned against excessively
heavy summer pruning as this may cause shock, resulting in an inhibition of
fruit growth and promotion of russeting. Lemmens (1982) pointed out that
severe summer pruning may lead to small and poorly coloured fruits. Nelgen
(1980) reported a reduced relative fruit weight and more bitter pit in severely
vs. moderately pruned trees.

3.3. E n v i r o n m e n t

Gardner et al. (1952) stated that environmental conditions, particularly

nutrient supply, soil moisture and light, greatly influence the nature of response
266

to summer pruning, and many contradictions between different reports can be

explained in this way. For instance, Marini and Barden (1982d) measured a
significant suppression of fruit diameter 3 weeks after pruning 'Delicious' trees
during the 1979 season, but there was no influence on fruit size in 1980. Taylor
and Ferree (1984a) suggested that the effect of summer pruning on fruit size
was mitigated by crop position, and they observed varying effects on fruit col-
our in different years. However, only a few reports explicitely relate variations
in external conditions to variations in pruning effects. Gardner et al. (1952)
reported that atmospheric conditions favouring a high transpiration rate pro-
moted fruit spur and flower bud formation with certain summer-pruning treat-
ments. They further warned that promoting shoot growth by summer pruning
in order to hasten the development of young fruit trees is advisable only where
the growing season is long enough to permit proper maturity of the secondary
shoots. In support of this, Stik5 and Laurinen (1982) did not recommend sum-
mer pruning under Finnish conditions because of the short growing season
there, which interferes with the delayed hardening of the trees after pruning.
When soil conditions, particularly moisture and nutrient supply, encourage
new vegetative growth, summer pruning is less likely to promote fruit spur and
flower bud formation than when less moisture and less nitrogen are available
(Gardner et al., 1952 ). Miller (1982) confirmed that the amount of regrowth,
as well as flower bud initiation on regrowth from stubs, varied with moisture
conditions during 2 growing seasons.

3.4. T r e e c h a r a c t e r i s t i c s

C u l t i v a r . ~ There are several observations that cultivars respond differently

to summer pruning with respect to growth, flowering or certain fruit charac-
teristics. However, there is no general understanding about why they respond
differently. Regarding vegetative growth, the final shoot length is influenced
by the vigour of the cultivar (Mika et al., 1983). Aselage and Carlson (1977)
noted that each variety responds differently to summer pruning treatments,
and the proper timing of pruning, in particular, very much depends on the
cultivar concerned. Marini and Barden (1982d) pointed out that summer
pruning is advisable mainly in cultivars suffering from poor colouration, but it
may contribute little to the colouration of high-colouring strains.

Vigour. w Vigour of apple trees does not depend only on environmental factors
(cf. Section 3.3 ) or on cultivar, but can also be influenced by rootstock, crop-
load and age. Bos (1974) reported that the inhibition of growth by summer
pruning occurred only on M 9, and not on more vigorous rootstocks. Mika et
al. (1983) noted that the pruning response increased with the vigour of the
scion and rootstock, although the scion/rootstock influence was less pro-
nounced aftersummer pruning than afterdormant pruning. Myers and Ferree
 267

(1983b) suggested that differences in scion/rootstock combinations may

explain many of the differences and contradictory conclusions in summer
pruning research. In trees with a full crop load, there was a distinctly greater
reduction of stem thickening caused by summer pruning than in non-bearing
trees, according to experiments by Taylor and Ferree (1984a).

4. THE PHYSIOLOGYOF PLANT RESPONSES

Shoots which are removed by summer pruning while plants are non-dor-
mant, or in the state of predormancy (Saure, 1985b), have leaves, buds and
often growing shoot tips. However, most of the diverse reponses to summer
pruning in growth, flowering and fruit development, and most of the theoret-
ical conclusions drawn from these summer pruning effects, have been attrib-
uted to the loss of foliage only.

4.1. Removal of leaves

There are mainly two aspects of leaf removal that have been considered in
relation to summer pruning effects:
- - loss of carbohydrate supply due to the reduction of photosynthetically active
leaf surface;
better penetration of light to the interior of the canopy.

Reduced carbohydrate supply. - - Gardner et al. (1952} suggested that the

immediate effect of any summer pruning on the tree as a whole is to reduce the
carbohydrate supply and the rate of carbohydrate production, and to increase
the supply of water and nutrients available to the rest of the plant. They spec-
ulated that under certain conditions this may cause a depletion of the reserves
for the following season, and that growth may be correspondingly restricted in
that year. They further suggested that where photosynthetic processes are seri-
ously reduced, fruit quality would certainly be impaired.
In apple fruits, a reduction of soluble solids levels has been confirmed by
most authors (cf. Section 2.4). H. Prigge (personal communication, 1981)
observed that such fruits were damaged on the tree by early winter freezes,
whereas those from unpruned trees were not. Similarly, a reduction of carbo-
hydrates throughout the winter has been observed by Maggs (1965) in young,
non-fruiting apple trees pruned in August. Their stems contained no starch
and their late-formed wood was not lignified. Priestley (1964) found the amount
of new growth of young apple trees did not depend on the amount of initial
reserves, but Head (1969) and Kandiah (1979) maintained that the effect of
reserves cannot be excluded completely. Engel and Lenz (1981) observed that
summer pruning in September, removing 60% of the leaves after termination
of growth, caused a slightly lower starch and sucrose content of the shoots and
268

buds, and the resistance of the flowers to late spring frostswas severely reduced
in the next season compared to winter pruned trees. However, the lower frost
resistance was not closely correlated to the lower carbohydrate content. Even
complete defoliation, resulting in a great loss of starch and sucrose in shoots
and buds, produced fewer frost-damaged flowers than the summer pruning,
and partial defoliation (44%) of all current season's shoots did not alter the
frost resistance of the flowers.
Taylor and Ferree (1981) and Myers and Ferree (1983a) reported that sum-
mer pruning reduced the relative amount of dry matter accumulation in roots,
as indicated by lower values for the dry-weight root/shoot ratio. Complete
defoliation of young apple trees 4-6 weeks before natural leaf-fall greatly
reduced root growth within a few weeks of treatment (Head, 1969). This is in
agreement with earlier findings of Richardson (1957) with Acer saccharinum,
in which defoliation of seedlings brought about a cessation of root elongation.
However, Taylor and Ferree (1986) stated that 11 weeks after summer pruning
the concentrations of gross water-soluble reducing sugars and insoluble hydro-
lyzable carbohydrates in the roots was not significantly affected by that prun-
ing; those of sucrose and fructose even increased with increasing severity of
summer pruning.
Maggs (1964) observed that partial defoliation gave rise to a compensatory
increase in photosynthetic activity in the remaining leaves. Taylor and Ferree
(1981) also reported that after summer pruning net photosynthesis and the
transpiration increased substantially, especially in older leaves, with the extent
depending on the severity of pruning. The differences were noticeable even
over 1 month after pruning. These findings have been confirmed by Marini
and Barden ( 1982c ) and Myers and Ferree (1983a), who stressed that summer
pruning effects on photosynthesis resulted from a delayed decline in photosyn-
thesis as compared to unpruned controls. They agree with results in other crops
such as mulberry (Morus alba) (Satoh et al., 1977) and peach (Rom and
Ferree, 1985).
Taylor (1982) concluded that summer pruning effects on shoot elongation,
fruit growth and root growth do not appear to be controlled by carbohydrate
levels or type of carbohydrate present. Gardner et al. (1952) reported that
certain parts of the plants can be favoured by summer pruning, but other parts
may suffer. De Haas and Hein (1973) noted an almost complete cessation of
root growth at the same time as new secondary shoots were being formed after
summer pruning. These findings may be taken as an indication that there could
be a redistribution of assimilates to the pruned parts of the plant rather than
an absolute deficit. This has been further investigated by Mika and Anto-
szewski (1973), Marini and Barden (1983) and Rom and Ferree (1985). This
redistribution is probably controlled by the buds rather than the leaves (Naga-
rajah, 1975 ), as will be discussed in Section 4.2.
 269

Improved light penetration, a Improved light penetration for better fruit col-
ouration is one of the main objectives of summer pruning. It is usually done
rather late in the season in order to limit adverse effects on fruit development
(cf. Section 2.4). Gardner et al. (1952) pointed out that early pruning, espe-
cially when heading cuts are used, may cause the formation of many secondary
shoots and result in heavier shading of the leaves and fruits in the lower parts
of the tree. A highly positive correlation between percentage of red blush and
photosynthetic photon flux density has been established by Morgan et al.
(1984) after early ( 73 days after full bloom ) and late (108 days after full bloom
or 4 weeks before first harvest) pruning, but more importantly, the increase
in red blush was relatively greater in those heavily shaded zones of the trees
where light transmission was not markedly increased by summer pruning and
where, independently of the pruning treatment, the mean concentration of
soluble solids was lowest. No explanation for this observation has been offered.
Gardner et al. (1952) noted that moderate early summer pruning, preferably
done by thinning, allows more light to reach leaves on the lower parts of the
shoots, and therefore tends to encourage fruit spur and flower bud formation.
This view has been supported by many authors since, but there is no experi-
mental evidence that such an effect of summer pruning is caused specifically
by better light penetration. Porpiglia and Barden {1981) could confirm that
the diffuse photosynthetically active radiation increased immediately after
summer pruning, with the percentage increase depending on whether it was
measured in the periphery of the tree or within the canopy, but the rates of net
photosynthetic potential and dark respiration of interior spur leaves remained
consistently lower 3 weeks after summer pruning than those of leaves from the
periphery. Actually, they did not differ significantly from leaves of a similar
location on unpruned trees. Heading even reduced light penetration through-
out the canopy the year after treatment, due to a proliferation of shoots devel-
oping immediately below the heading cuts. Marini and Barden (1982b) also
measured an increase in light penetration throughout the tree canopy for the
remainder of the season after summer pruning. The specific leaf weight of
peripheral leaves did not decline following summer pruning, in contrast to the
decline in dormant-pruned trees. The response was probably due to improved
light conditions and/or delayed leaf senescence. In the next season, light pen-
etration was reduced and the specific leaf weight of interior leaves was lower
than that of the interior leaves of dormant-pruned trees.
As well as allowing better light penetration, summer pruning could improve
the penetration of pesticides and other sprays. Bangerth and Link (1972)
assumed that summer pruning was partially effective in reducing the incidence
of bitter pit by enabling better deposition of Ca sprays in the tree interior.

4.2. Removal of buds and shoot tips

The effect of removing buds and growing shoot tips has rarely been discussed
when trying to understand plant responses to summer pruning. However,
270

growing shoot tips are known to be important sources of auxins which play a
central role in apical dominance (Phillips, 1975 ). Saute (1971) has postulated
that auxins have a key role in the control of growth and developmentby acting
as controllers in the cybernetic system based on the interaction of shoots and
roots. Via their influence on root activity, they may cause a positive or negative
feedback, depending on their concentration. Accordingly, pruning would inter-
fere with this natural regulation by removing active or potential sites of auxin
production. This hypothesis has been outlined comprehensively elsewhere
(Saure, 1981 ).

Adjacent leaves, and subsequently grow-

Release [rom correlative inhibition. - -
ing shoot tips and terminal buds, are known to prevent the breaking of lateral
buds. This correlative inhibition is one aspect of apical dominance. Removing
important sites of auxin production by summer pruning, except if done by
thinning cuts, temporarily eliminates the inhibition imposed on the lateral
buds, and consequently promotes their outgrowth. However, this promotion of
axiUary bud growth can only be achieved when the buds are still under the
external control of leaves or shoot tips. Later in the season, when they have
reached the state of true dormancy which is under the control of factors inside
the buds, pruning is ineffective in causing bud break (Saute, 1985b).

Redistribution of nutrients, m The other component of apical dominance is the

sink activity exerted by sites of high auxin activity. Apical dominance causes
the preferential movement of nutrients, and probably of phytohormones like
cytokinins (Lockard and Schneider, 1981), to the growing shoot tips, espe-
cially in non-fruiting trees. In fruiting trees the fruits, by their own auxin pro-
duction, have even greater capacity to attract nutrients than shoot tips. Thus
fruits are strong competitors m and are at least as strong as growing shoot tips
in their ability for correlative inhibition of the shoots below them. Therefore
removing the sites of auxin production not only removes correlative inhibition,
but may also cause a temporary redistribution of nutrients. Quinlan and Pres-
ton (1971) showed that repeated pinching of a growing bourse-shoot modified
the distribution pattern of assimilates. Apical photosynthate movement out of
a primary leaf towards the growing shoot tip was shifted by the pinching to
mainly basipetal transport towards the flower clusters and the fruitlets, which
is the usual direction of transport in bourse-shoots after termination of growth.
However, Mika and Antoszewski (1973) observed that after early (June)
pinching in young, vegetative apple trees, the absence of strongly competing
sinks meant that photosynthates accumulated mainly below the pinching point
and were hardly translocated to more distal parts of the plant.
Roots are considered to be poor competitors within a plant for photosyn-
thetic assimilates (Radin et al., 1978 ). Only after termination of shoot growth,
when the auxin activity of the shoot tips is lowered, will the nutrients be trans-
 271

ported increasingly to the roots (Hansen, 1967). The stabilizing mechanism

maintaining the balance of distribution between the various regions in young
trees, which Maggs (1965) was looking for, probably works in this way.
As indicated in Section 4.1, root development may be reduced by summer
pruning, but this reduction apparently starts only after new shoot growth has
started. The amount of reduction was related to the intensity of shoot growth
(Head, 1967). Consequently, inhibition of shoot regrowth may promote root
growth, as shown shortly after treating young apple trees with the growth
retardant Alar by Schumacher et al. (1971). In experiments by Polefska (1983),
tipping + partial disbudding ( no regrowth) resulted in the greatest dry matter
accumulation in the root system of young apple trees, 2 and 4 weeks after
treatment; pruning or defoliation, causing regrowth, reduced dry matter accu-
mulation as compared to unpruned controls. Marini and Barden {1983) noticed
that in young apple trees pruned in September after termination of growth,
photosynthate from basal leaves was primarily transported to regrowth 3 weeks
after pruning, but root-directed transport - - similar to unpruned trees and
immediately after pruning - - was maintained if regrowth was prevented by
NAA treatment. Experimental evidence that it is IAA that directs the trans-
port of photosynthate has been provided by Altman and Wareing (1975), who
applied IAA to the base of cuttings and caused basipetal transport of assimi-
lates and accumulation of sugar in this region. The increased transport caused
by auxin probably also accounts for the known induction of transport channels
(sieve tubes) by auxins ( Sachs, 1975).
However, the effects of summer pruning on root growth cannot be attributed
only to an altered nutrient supply caused by a modification of the sink capacity
of the shoots. From defoliation and disbudding experiments in young A c e r sac-
c h a r i n u m seedlings, Richardson (1957) concluded that hormones were involved
because defoliation led only to a cessation of root elongation, but disbudding
completely suppressed root formation.
A promotion of root development by leaves and buds seems to contradict the
inhibition of root growth by growing shoots, as mentioned above. However,
these differences may be taken in support of the hypothesis that the roots act
as some kind of correcting unit in a control loop in which growth can be con-
trolled by positive or negative feedback depending on the concentration of aux-
ins (Saure, 1971, 1981). This hypothesis is supported, for example, by the
observation of Pilet et al. (1979) that the inhibitory effect of exogenously
applied auxin on growth of root segments can be shifted to growth promotion
by reducing the endogenous auxin concentration. Further information could
be obtained by comparing the immediate effect of pinching in young, non-
dormant apple trees having few growing shoots with the effect in older trees,
but so far such root investigations are known for young trees only.

A c t i v a t i o n o f roots, w The promotion of regrowth by summer pruning is often

272

considered to be a resultof an altered root/shoot ratio,which implies a greater

concentration of root-produced growth promoters in remaining leaves (e.g.
Polefska, 1983 ). However, there is strong evidence of an initialroot activation
after eliminating the inhibition caused by the shoots. Skene (1968), who
observed a substantially increased cytokinin level in the bleeding sap of grapes
after growth was retarded by C C C treatment, was not sure whether this was
directly related to cytokinin synthesis by the roots, or to the larger root mer-
istem. Beever and Woolhouse (1974), who removed apical and lateral buds
from vegetative plants of Perilla frutescens, noted that a decreased rate of root
elongation was accompanied by an increased cytokinin export from the roots.
Certainly, an increased level of cytokinins in the uppermost remaining leaves
was recorded soon after pinching of mulberry shoots (Satoh et al., 1977), and
in sprouting grape buds after summer topping (Matsui et al., 1979). However,
it needs to be determined whether the cytokinins observed are always of root
origin, or whether they may also be transformed from storage forms which are
found in the bark and older leaves (Van Staden and Davey, 1979).

Retardation of senescence. ~ Senescence is a very important process, not only

in the life cycle of the trees but also in the annual cycle of development in apple
trees, as this phenomenon is associated, e.g., with:
E spur development (reduced apical dominance; Wareing, 1970);
accumulation of nutrients;
reproductive development (flower induction);
ripening;
abscission;
induction of dormancy;
frost hardiness.
Cytokinins are known to inhibit senescence, i.e. to prevent or even reverse
the senescence of leaves. The effects of cytokinin treatment, such as leaf
enlargement, re-greening with increased levels of chlorophyll, protein and RNA,
and consequently increased photosynthesis, are identical to those observed
after summer pruning in apple (Taylor and Ferree, 1981; Marini and Barden,
1982c; Myers and Ferree, 1983a; Ferree et al., 1984), peach (Rom and Ferree,
1985) and mulberry (Satoh et al., 1977). Satoh et al. (1977) stated that the
removal of axillary buds additionally to pinching aggravated the effects of shoot
pruning, e.g. with respect to increased photosynthesis, delayed starch increase,
maintained leaf chlorophyll, and continued mesophyll cell division. These
results suggest that an increase in free cytokinins after summer pruning seems
to be an important factor in pruning-responses of apple trees. Factors inhib-
iting the cytokinin export from the roots, e.g. root pruning in apple (Geisler
and Ferree, 1984), peach (Richards and Rowe, 1977) and other crops (McDavid
et al., 1973), or waterlogging (Childers and White, 1942) or drought, cause
opposite effects in that they promote leaf and plant senescence.
 273

Additionally, summer p r u n i n g - - if heading cuts or pinching are u s e d - - may

prevent or delay senescence in a more direct way also. It has been shown that
leaf senescence can be promoted by nearby growing regions, and the elimina-
tion of younger organs has consequently retarded or reverted senescence
(Nood~n and Leopold, 1978; Thomas and Stoddart, 1980). This agrees with
the proposed double function of growing shoot tips in controlling growth directly
by correlative inhibition, or indirectly by controlling root activity. In apple
trees, the rejuvenation caused by summer pruning was accompanied by a great
reduction of ABA activity in the buds within 6 days after pruning (Myers,
1981 ), and in mulberry trees the ABA level of the uppermost leaves remaining
after summer pruning was also substantially lowered within 6 days ( Satoh et
al., 1977).
Obviously, the rejuvenation caused by summer pruning does not result only
from the removal of buds and growing shoot tips. Taylor et al. (1984) observed
a reduction in the ABA activity in the apex tissues of closed terminal buds on
short shoots, and a marked increase in gibberellin-like substances, also after
defoliation, but then the increase in cytokinin-like substances was only slight.
The effect on ABA and GA content, and the amount of bud break, were more
marked when defoliation was done early, pointing to an advance in true dor-
mancy with later defoliation. However, from the observation of Engel and Lenz
(1981) on frost damage in apple blossom after various summer treatments (cf.
Section 4.1), we may conclude that the rejuvenation by summer pruning is
stronger than that caused by total defoliation, provided the assumption is right
that the reduced frost hardiness was primarily caused by disturbed onset of
dormancy.
It is not known whether the same mechanisms acting in leaf senescence are
also responsible for the delay in certain characteristics of fruit maturation (cf.
Section 2.4). In tomato (Lycopersicon lycopersicum), Varga and Bruinsma
(1974) observed that a reduction in foliage considerably increased the cyto-
kinin level in the fruits, and that ripening was progressively retarded by
increasing levels of endogenous cytokinins. This observation seems to explain
the effect of summer pruning on fruit maturation better than an explanation
based only on reduced carbohydrate supply following a reduction of supporting
leaves. It could also explain the increased ethylene levels, as observed by Tay-
lor and Ferree (1984b) in maturing apples of summer-pruned trees (cf. Sec-
tion 2.4). There are numerous reports indicating that ethylene production is
raised at higher cytokinin activity, but simultaneously its inhibiting effects are
prevented by cytokinins (Saute, 1985b). In support of the observations in
tomatoes, apple growers in Northern Germany have observed that early removal
of water sprouts may halt fruit development for several weeks (verbal com-
munication), although water sprouts consume rather than supply carbohy-
drates. Unfortunately, the effect of cultural practices on the content of
274

phytohormones in apple fruit, and the influence of the resulting hormonal

changes on physiological processes, still await investigation (Ebert and Ban-
gerth, 1985).
Cytokinins could also play a role in the predominantly positive effects of
summer pruning on anthocyanin synthesis which occurs even at low light lev-
els, and despite protracted dry matter increment and delayed maturation (cf.
Section 2.4). Besides increasing light penetration to the fruit, summer pruning
could increase sensitivity to light, i.e. an increased colouring capability could
exist, similar to that achieved by temporary covering of apple fruits during fruit
development (Kikuchi, 1964; Proctor and Lougheed, 1976). Cytokinin treat-
ments were found to cause an intensive red colouration under certain condi-
tions in young shoots of conifers (Kossuth, 1978; Mulgrew and Williams, 1985).
However, this aspect of colour promotion has been widely neglected so far and
still requires a lot of research.
The information available for discussion in this section is complex and not
well elucidated, and the picture drafted of the physiology of summer pruning
effects is far from being complete. For instance, it is well established that aux-
ins exert their influence on senescence and ripening at least partially by their
influence on, and their interaction with, ethylene. Further, it is known that
the primary effects of cytokinins are accompanied by variations in the biosyn-
thesis and/or activity of gibberellins (Grochowska et al., 1984 ). However, the
role of gibberellins in rejuvenation, and their interaction with cytokinins, are
still insufficiently understood. Yet, even the limited information presented here
will be sufficient to indicate the necessity for re-evaluation of some established
theories regarding the physiology of summer pruning effects for the benefit of
commercial fruit production.

5. SUMMERPRUNING AS AN INTEGRATEDCULTURALPRACTICE

As shown above, summer pruning, like pruning in general, is a cultural prac-

tice altering the capacity of the trees to control their growth. The endogenous
regulation usually becomes active when growth exceeds a certain threshold. If
this balance is disturbed, plants tend to re-establish it by increased growth/re-
growth ("parachute effect"; Saure, 1981). Therefore, both summer pruning
and dormant pruning must be considered primarily as growth-promoting prac-
tices, but the invigoration is rather short-term. Accordingly, Maggs (1965)
pointed out that dormant and summer pruning are not such diametrically
opposed treatments as issometimes suggested, and other authors, e.g. Prigge
(1981), Marini and Barden (1982c) and Saute (1985a), confirmed that the
effects of both treatments on growth vigour are similar.
The fundamental difference between them is that summer pruning is done
while the trees are non-dormant or in the state of pre-dormancy, which is at
least partially reversible. Therefore, summer pruning delays all phenomena
related to senescence (cf. Section 4.2 ). Dormant pruning deals with trees that
 275

have completed this process, or have already overcome true dormancy rather
recently. Therefore, while dormant pruning can only influence the source/sink
ratio still to be established in the coming season, summer pruning a d d i t i o n a l l y
interferes with an active shoot/shoot, shoot/root, or shoot/fruit interaction.

G r o w t h r e d u c t i o n . ~ It may be concluded that growth of apple trees with a

well-functioning root system generally cannot be reduced by any kind of prun-
ing (this is probably different for newly planted trees). Therefore, although
pruning is the only way to reduce tree size, and especially the unproductive
interior of the canopy, it should be avoided where shoot vigour is to be reduced.
The growth promotion and - - in the case of summer pruning - - the delaying
effect on dormancy are more pronounced the more invigorating factors prevail
such as fertile soils, warm and humid climate, vigorous rootstocks, juvenility
of the plant, and poor fruit set. The growth responses will also increase with
the severity of pruning, and will be stronger after heading or stubbing than
after thinning. If it is necessary for other reasons, any kind of pruning should
be accompanied by weakening cultural practices such as bending of shoots,
reduced N supply, or reduced irrigation.

Flower bud formation. -- To influence flower bud formation, the timing of

pruning is decisive. Flower bud formation may be taken in principle as a 3-step
process, consisting of:
(1) the development of vegetative meristems in the bud initials;
(2) the conversion of these meristems into generative meristems;
( 3 ) the further development of these generative meristems.
Consequently, early summer pruning may be expected to benefit flower initi-
ation in the apical buds of spurs and short shoots only if tree vigour is poor, by
promoting Step 1. Late summer pruning may then promote the further devel-
opment of these flower initials ( Step 3) and, if thinning cuts are used, may
encourage flower initiation in the lateral buds of current season's shoots. How-
ever, in vigorous trees, early summer pruning can be expected to disturb flower
initiation in the apical buds by preventing Step 2, which requires a temporary
inhibition of bud growth. This negative effect of early pruning in vigorous trees
will be aggravated by increasing severity of pruning, by heading and stubbing
instead of shoot thinning, and by accompanying invigorating factors. In lateral
buds, repeated pinching may encourage flower bud formation by promoting
Step 1.

Fruit development. ~ In vigorous trees, the eating quality of fruits is likely to

be impaired most by early and severe summer pruning, especially when the
growing season is short. The risk of June-drop will increase in these trees cor-
respondingly. Repeated early pinching may reduce June-drop, but would be
very laborious. Light summer pruning, preferably by thinning, may help to
276

improve fruit quality only where external conditions prevent vigorous growth.
In the season after summer pruning, larger fruit can be expected provided that
flower bud formation has been reduced in the previous season, but this would
be at the expense of yield. The necessity of heavy late summer pruning for
better fruit colour ("exposure pruning") may be taken as an indication of
undesirable tree development due to natural growth factors and/or inappro-
priate cultural practices that need to be corrected, but better should have been
avoided.

ACKNOWLEDGEMENTS

Dagmar Geisler-Taylor and Dr. B.H. Taylor have been very helpful in crit-
ically reading the manuscript, and I am very grateful for their suggestions. I
thank all those who have provided me with literature, and who have encour-
aged me to prepare this review.

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