New Zealand Journal of Crop and Horticultural Science

ISSN: 0114-0671 (Print) 1175-8783 (Online) Journal homepage: http://www.tandfonline.com/loi/tnzc20

Competition between fruit and vegetative growth

in Hayward kiwifruit

PEH Minchin , WP Snelgar , P Blattmann & A J Hall

To cite this article: PEH Minchin , WP Snelgar , P Blattmann & AJ Hall (2010) Competition
between fruit and vegetative growth in Hayward kiwifruit, New Zealand Journal of Crop and
Horticultural Science, 38:2, 101-112, DOI: 10.1080/01140671003781728

To link to this article: https://doi.org/10.1080/01140671003781728

Published online: 01 Jun 2010.

Submit your article to this journal

Article views: 740

Citing articles: 10 View citing articles

Full Terms & Conditions of access and use can be found at

http://www.tandfonline.com/action/journalInformation?journalCode=tnzc20
New Zealand Journal of Crop and Horticultural Science
Vol. 38, No. 2, June 2010, 101
112

Competition between fruit and vegetative growth in Hayward kiwifruit

PEH Minchina*, WP Snelgara, P Blattmanna and AJ Hallb
a
The New Zealand Institute for Plant & Food Research Ltd, Plant & Food Research Te Puke, 412 No 1 Rd,
RD2, Te Puke 3182, New Zealand; bPlant & Food Research Palmerston North, Private Bag 11600, Manawatu
Mail Centre, Palmerston North 4442, New Zealand
(Received 10 August 2009; final version received 26 February 2010)

Fruit dry matter content (DM) of kiwifruit, defined as the fruit dry weight (DW) expressed as a
percentage of the fresh weight (FW), is used as an indicator of eating quality. A high DM at
harvest results in increased consumer satisfaction. To determine the role of carbohydrate
availability and sink competition in altering fruit size and DM at commercial harvest, we used an
experimental model system based on phloem-girdled shoots to manipulate the number of source
leaves, fruit and vegetative sinks on a shoot of ‘Hayward’ kiwifruit. Commercially produced
kiwifruit fruits typically weigh 95
115 g and have a DM of 14
17 units (this is a percentage
but the % sign has not been shown, to reduce confusion with changes in DM expressed as a %
change). With our experimental system, this corresponded to the fruit growth observed with two
to three leaves when there was no vegetative competition. With four source leaves and a single
fruit, competition from a vegetative regrowth reduced fruit fresh weight by 28% and dry weight
by 39%. With two fruit, and no vegetative sink, between-fruit competition had less severe effects
on the fruit and dry weight. Doubling the number of fruit on a girdled shoot reduced average
fruit fresh weight by 15%, dry weight by 23% and DM by 12%. With a high source supply
(seven leaves with no regrowth competition) fruit growth was very high with a fresh weight
of 174 g and 18.4 DM, substantially above that of commercial production.
Keywords: Actinidia; vegetative competition; fruit growth

Introduction often stated as a percentage. In this work we do

Consumers prefer kiwifruit that are large
and have a high soluble solids content (SSC)
when eating-ripe (Crisosto & Crisosto 2001;
Lancaster 2002; Harker et al. 2009). Since
kiwifruit are picked before the starch in the
fruit has been fully converted to soluble sugars,
it is not possible to measure the eating-ripe
SSC at harvest. However, dry matter (DM)
measured as dry weight (DW) or measured as a
percentage of total fresh weight (FW) concen-
tration of a fruit, provides a good estimate of
ripe SSC (Richardson et al. 1997; Jordan et al.
2000). DW and FW are measured in grams,
and DM is expressed as the percentage of the
FW that is DW, that is 100 DW/FW which is
not express DM with units of % as this can lead
to confusion when comparing differences in
treatment effects that are often expressed as a
percentage change, so the two uses of percen-
tage have the potential to be confused. In this
work when percentage (%) is used, it refers to
a treatment change expressed relative to the
control. New Zealand kiwifruit orchardists
are now paid a premium for fruit that meets
the DM requirements of customers (Woodward
et al. 2007). In commercial orchards, kiwifruit
typically have a fresh weight (FW) of 95
115 g
(Taylor et al. 2007) and DM of 14
17 (Burdon
et al. 2004).

*Corresponding author. Email: peter.minchin@plantandfood.co.nz

ISSN 0114-0671 print/ISSN 1175-8783 online
# 2010 The Royal Society of New Zealand
DOI: 10.1080/01140671003781728
http://www.informaworld.com
102 PEH Minchin et al.
Kiwifruit vines produce excessive vegetative
growth which competes with fruit growth for
carbohydrate and other resources as well as
creating shade. Modern canopy management
methods are aimed at reducing vegetative
growth and, over the past 15 years, these
techniques have almost doubled the yield of
high-quality fruit per vine. The physiological
principles behind these changes are not well
understood, so we do not know if we have
reached the limit of improvements in harvest
yield, or DM production.
Dry matter predominantly enters the fruit
through the phloem while water enters via both
the phloem and xylem and is lost by fruit
transpiration. Hence DW is a measure of
accumulated phloem flow, whereas DM is a
complex mix of phloem, xylem and transpira-
tion flows. An ecophysiological model of fruit
growth and development of peach driven by
phloem, xylem and transpiration flows has
successfully reproduced important fruit traits
including fruit FW and DW (Fishman &
Génard 1998; Génard et al. 1998; Lescourret
et al. 1998, Lescourret & Génard 2005) and
recently has been used to describe genotypic
variation within breeding populations (Quilot
et al. 2005).
Higher plants consist of many sources of
carbohydrate and many carbohydrate sinks.
Growing plants orchestrate the balance
between supply and demand to develop and
mature in a way characteristic of the species. A
single sink is prevented from dominating. In the
case of more distributed reproductive sinks,
there is more variability in fruit size, but this
is still kept under remarkable control (Avery
et al. 1979; Palmer 1992; Austin et al. 1999).
Maximum fruit size is currently believed to be
limited by its growth potential determined early
in development, probably during an early stage
of cell division (Grossman & DeJong 1995).
Carbohydrate sinks are described by their sink
strength, that is their ability to attract carbohy-
drate when supply is not limited (Waring &
Patrick 1975; Grossman & DeJong 1994;
Lacointe & Minchin 2008), and by their
priority which describes their ability to compete
when carbohydrate supply is limited. In the
species that have been investigated to this
detail, sink priority increases from roots to
reproductive sinks in the following order
(Wright 1989; Wardlaw 1990):
storageBroots Bcambium; fleshy fruit parts;
shoot apices; leaves Bseeds
A high priority sink, for example the seeds,
can have a small total requirement but when
supply is reduced, it is the high priority sinks
that get the limited resources. In contrast, roots
have a low priority, such that at times of short
supply they are starved of carbohydrate.
Experiments with apple trees have shown that
roots can die from lack of carbohydrate supply
when they are over cropped (Palmer 1992).
The aim of this work was to quantify the
effects of competition from vegetative sinks
upon FW and DW accumulation with the fruit
of kiwifruit vines. This information is needed
for mechanistic understanding of these interac-
tions and to determine what controls fruit DM.
Materials and methods
Experimental sites and treatments
Experimental work was carried out on the
Te Puke Research Orchard, Bay of Plenty,
New Zealand (37849?S, 176819?E) during
the 2006
08 seasons. Mature (10 year-old)
Actinidia deliciosa (A. Chev.) C.F. Liang et
A.R. Ferguson ‘Hayward’ kiwifruit trained on
a pergola trellis were used.
The trial work was carried out on vines
with an open canopy and with vigorous growth
to avoid weak self-terminating shoots. This
requirement meant that a different orchard
block of kiwifruit vines was used each year.
In year 1, the selected block had been managed
to organic BioGro standards (BioGro 2009). In
year 2, a different orchard block was used, with
vines managed conventionally, which included
use of hydrogen cyanamide to increase bud
break and flowering.
Flowering was monitored on 20 tagged
canes in the experimental block. The number
of fully open flowers was assessed at about five-
day intervals and the date of mid-bloom
estimated as the 50% point of a fitted cumu-
lative curve of the total number of open flowers
verses date. Fruit growth is referenced as days
after mid-bloom (DAMB).
 Competition between fruit and vegetative growth in Hayward kiwifruit 103

When a growing kiwifruit shoot is pruned,
the remaining most distal bud will usually
break and develop into a secondary shoot,
which is also known as a regrowth. We used
shoot pruning as a method to induce a vegeta-
tive sink located near the fruiting zone to
compete with a developing fruit.
Experimental shoots were initially pruned
to length so that they contained seven leaves
and two to four fruit. These leaves were mature
and quite large (c. 130 cm2, diameter c. 13.0 cm
[Snelgar & Thorp 1988]). All shoots were
phloem girdled to restrict the flow of carbohy-
drate in or out of the shoot, and this girdle was
kept open until fruit were harvested. In both
years, shoots were pruned 10 days after flower-
ing, thus stimulating regrowth. Girdles and the
treatments were applied 10 days after that. In
year 1, all unwanted regrowths were removed
at about 50 DAMB and in year 2 at about 20
DAMB. An example of a girdled experimental
shoot with four leaves, one fruit and a regrowth
is shown in Fig. 1.
Each treatment was replicated 20 times, and
all treatments were applied in a randomized
block design. Sink/source relationships were
manipulated by:
1. Adjusting source potential by setting the
number of mature leaves on each shoot
to two, four or seven. These are denoted
2L, 4L and 7L. Previous work has
indicated that with six or more leaves,
supply to the fruit was near saturation
(Snelgar et al. 1986). We designed these
experiments with most fruit supported
by four leaves so as not to saturate
growth of a single fruit. Under these
source-limited conditions, any compet-
ing sink would be expected to signifi-
cantly reduce fruit growth. In year 2, all
shoots were pruned to four leaves. The
number of fruit on the shoot is denoted
by 1F or 2F. In year 1, only the 1F
treatment was used; in year 2, both 1F
and 2F treatments were used.
2. A vegetative sink was added by allowing
a single vegetative regrowth to develop
from the distal end of the primary shoot
after pruning (see Fig. 1). Since growth
rates of vegetative shoots are typically
50 mm/day (Snelgar et al. 2005), these
were expected to be strong sinks. To
maintain the regrowth as a carbohydrate
sink, leaves were removed once they
reached about 50 mm in diameter. At
this early stage of growth, the leaves were
expected to be net sinks (Lai et al. 1988;
Greer & Jeffares 1998; Greer 1999).
These defoliated regrowths are denoted
R. On all other shoots, where no vege-
tative sink was wanted, regrowths were
removed before they were 100 mm long.
3. In year 2, treatments involving four
leaves with one fruit in competition

Fig. 1 A schematic diagram of the experimental unit for treatment 4L1FR showing the girdles at the base of
the shoot. Initially the shoot has been pruned to length so that seven leaves and two fruits remained, and
girdled to phloem isolate the experimental unit. About 20 days later, this was reduced to four leaves and one
fruit. The initial pruning stimulated development of a vegetative regrowth and this was retained. Leaves were
continually removed from the regrowth before they were fully expanded. In this way the regrowth acted as
a net sink.
104 PEH Minchin et al.
with a de-leafed regrowth were repeated.
In addition, treatments with two com-
peting fruit were added.
In year 1, the full set of treatments was
2L1F, 2LR, 4L1F, 4LR, 4L1FR, 7L1F, 7LR.
In year 2 we repeated treatments 4L1F, 4L1FR,
4LR and added 4L2F.
Monitoring
Fruit growth was monitored non-destructively
by measuring the fruit length (L) and the
maximum (D1) and minimum (D2) fruit dia-
meters in mm then calculating the product
D1 D2 L/1000, which is denoted as LDD.
LDD was converted into an estimate of the
fruit FW using a calibration curve derived from
destructive fruit harvests made at various times
including fruit harvest (Snelgar et al. 1992).
Non-destructive measurements were made at
10
30 day intervals throughout the growing
season. A non-destructive measurement of
regrowth was obtained by monitoring regrowth
length.
Destructive sampling
In year 1, 20 samples of each treatment were
harvested at 50 and 100 DAMB and at fruit
maturity (about 180 DAMB). Linear dimen-
sions of the fruit were used to obtain a relation-
ship between LDD and fruit FW. Destructive
measurements of FW and DW of the fruit and
shoots were made. In addition, at 181 DAFB
(days after full bloom) seed counts were made
after extraction of the seeds from fruit softened
with pectinase (Rohapect† D5L Special, AB
Enzymes GmbH, Darmstadt, Germany). In
year 2, destructive measurements were made
at 180 DAMB.
Data analysis
The fruit growth rates were estimated by spline
fitting to the FW data using R software. This
method was used as it involves smoothing of
the observed data which reduces noise in
gradient estimation. Simpler differencing
of data increased observation noise but did
produce the same general patterns (not shown).
Means and their standard error were calcu-
lated using Microsoft Excel. Payton et al.
(2003) have shown that when the standard
errors do not differ by more than a factor of
two, the probability of overlap of 85.6%
confidence intervals is 0.95, that is, non-overlap
indicates p B0.05. A 85.6% confidence interval
corresponds to91.46 se. So in this work we
have used the rule that when the 1.5 standard
errors do not overlap, the means are significantly
different at p B0.05. As we do comparisons
within table columns as well as between data in
different tables, we have chosen not to indicate
non-significant differences within the tables,
leaving the reader to compare the means91.5
se which is a simple mental exercise and can be
done on any pair of means taken from the same
or different tables.
Results
Calibration of non-destructive fruit
measurements
Snelgar et al. (1992) demonstrated that the
kiwifruit fresh weight is highly correlated with
the production LDD of the three linear
measurements of a fruit, and is best described
by a power law. This work confirmed this, but
with slightly different parameter values. There
was very little difference between the power law
and a simpler linear relationship (Minchin et al.
2003), so we used the linear relationship in this
work, which was:
FW(g)0:658LDD r2 (adj)0:961
There was no indication of treatment differ-
ences, even though some treatments included
regrowth competition.
Fruit and shoot growth patterns
Table 1 lists the fruit data for year 1 measured
at harvest (181 DAMB). Reducing the number
of source leaves tended to increase variability in
both FW and DW. FW, DW and DM all
reduced with declining leaf number. Four
leaves were not sufficient to saturate growth.
In the presence of a regrowth, i.e. a vegetative
sink, the fruit FW, DW and DM were all
significantly reduced.
 Competition between fruit and vegetative growth in Hayward kiwifruit 105
Table 1 Year 1. Effect of leaf number (L) and regrowth (R) on fruit growth, measured as fresh weight (FW),
dry weight (DW) and dry matter content (DM), and on seed production, with standard errors in parenthesis.
Means within a column are different at a0.05, except for seed numbers which are not different at this level.
Each treatment shoot contained one fruit (1F)

Treatment FW (g) DW (g) DM Seeds

2L1F 103 (6.8) 15.6 (1.6) 14.7 (3.2) 1137 (95)

4L1F 153 (6.0) 26.5 (1.4) 17.3 (0.3) 1158 (49)
7L1F 175 (4.2) 32.3 (0.9) 18.4 (0.8) 1055 (74)
4L1FR 112.3 (4.9) 16.9 (0.9) 14.9 (0.3) 1080 (81)

Fig. 2A shows the time sequence of fruit
FW with the estimated rates of fruit growth.
The 4L1FR growth rate clearly shows two
rapid growth phases, with the second starting
at about 80 DAMB. It is usual to describe
the first and second rapid growth phases as
stage 1 and stage 3 with the intervening phase
as stage 2 (Coombe 1976). This is also clear in
the 4L1F data and possibly still evident in the
7L1F data but there is no sign of this multi-
phase behaviour in the 2L1F.
Fig. 2B shows the time sequence of
regrowth lengths. Regrowth was completed by
about 100 DAMB with no sign of a second
growth phase. Regrowth length increased when
more leaves were available to support growth,
as seen for fruit FW. With two leaves, regrowth
length was the least and with seven leaves the

200 1.6
2L1F A
180 4L1F
4L1FR 1.4
160 7L1F
estimated fruit FW(g)

1.2
140
growth rate (g/d)

1.0
120

100 0.8

80 0.6
60
0.4
40
rates 0.2
20

0 0.0
250
B
200
regrowth length (cm)

150

100

2LR
50 4L1FR
4LR
7LR
0
40 60 80 100 120 140 160 180 200
DAMB

Fig. 2 Time sequence of fruit and vegetative growth for treatments with varying numbers of leaves (L) and
fruit (F) in year 1. (A) Estimated fruit fresh weights (FW) based upon LDD. Lines with data points and
standard errors show the fruit FW, while the smooth lines alone are the corresponding growth rates
calculated using cubic spline fitting to the size data using the R statistical language. (B) Measured regrowth
lengths.
106 PEH Minchin et al.
200 40

fruit regrowth
150
30
fruit

DW(g)
FW(g)

100
20
regrowth
50

10
1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8
Number of leaves Number of leaves

Fig. 3 Effect of leaf number on fresh weight (FW) and dry weight (DW) at 181 DAMB of a single type
of sink (fruit filled symbols, regrowth hollow symbols). The single data point for four leaves when
there was also a sink of the other type competing for resources is shown as a triangle. Error bars show the
standard error.

maximum. The presence of a fruit reduced
regrowth length compared with that with the
same number of leaves and with no competing
fruit. Regrowth length with four leaves plus
fruit was greater than with two leaves and a
regrowth alone. Hence, the extra two leaves
were enough to compensate for the presence of
the fruit.
The effects of the number of source leaves
on fruit and shoot FW and DW at 181 DAMB
are presented in Fig. 3. Fruit FW and DW
increase non-linearly with leaf number, increas-
ing more rapidly with low leaf number and
showing clear signs of approaching saturation
with high leaf number. Regrowth FW and
DW increases linearly with leaf number, with
no indication of saturation. Regrowth DW
increases at a similar rate to that of fruit
DW for low leaf number and much more
rapidly as leaf number increases.
The FW and DW of fruit and vegetative
shoots for year 1, from destructive harvests at
100 and 181 DAMB, are presented in Table 2.
By 100 DAMB, a single fruit supplied by four
leaves (4L1F) had accumulated 109.5 g FW and
16.2 g DW. During the same period, a regrowth
supplied by the same number of leaves (treat-
ment 4LR) accumulated less FW and less DW.
The same was true for the entire 181 days.
Thus, for both periods, the accumulation in
both FW and DW of a vegetative sink alone
was less than that of a fruit alone.
Competition
With competition between the two sinks, in the
first 100 days, fruit FW was reduced by 23%

Table 2 Year 1. Average fruit and regrowth fresh weight (FW), dry weight (DW) and dry matter content
(DM), with standard errors (se) in parenthesis, for treatments with four leaves and only a fruit sink (4L1F),
only a vegetative regrowth sink (4LR), or both fruit and regrowth sinks (4L1FR). Means differ significantly
(PB0.05) when the difference is more than 1.5 times the sum of the standard errors (Payton et al. 2003)

Fruit Regrowth

Harvest Treatment FW (g) DW (g) DM (%) FW (g) DW (g) DM (%)

100 DAMB 4L1F 109.5 (3.0) 16.2 (0.5) 14.7 (0.2)

4L1FR 84.8 (7.9) 7.7 (1.1) 12.0 (0.4) 55.4 (4.0) 11.8 (0.8) 7.5 (0.7)
4LR 65.0 (3.4) 13.8 (0.8) 9.4 (1.3)
181 DAMB 4L1F 152.7 (6.0) 26.5 (1.4) 17.3 (0.3)
4L1FR 112.8 (4.9) 16.9 (0.9) 14.9 (0.3) 56.9 (3.4) 17.2 (1.1) 30.2 (0.7)
4LR 74.6 (7.2) 22.3 (2.2) 30.0 (0.6)
 Competition between fruit and vegetative growth in Hayward kiwifruit 107

and DW reduced by 52%, compared with fruit
alone. Regrowth FW was reduced by 15%,
DW by 14%, compared with regrowth alone,
with neither of these differences being statisti-
cally significant. Competition induced greater
FW and DW loss in the fruit than in the
regrowth. With competing fruit and regrowth,
total FW and DW growth was considerably
more than with either sinks alone, but less
than that of the fruit alone plus regrowth alone.
In the competing system, from 100 DAMB
to harvest, fruit FW and DW continued to
increase with DM also increasing. Regrowth
FW growth ceased, while DW continued to
increase, resulting in a large regrowth DM at
181 DAMB.
At 181 DAMB, vegetative competition
reduced fruit FW from 152.7 to 112.8 g, a
loss of 26% compared with a fruit alone. Fruit
DW was reduced from 26.5 to 16.9 g, a loss of
36% compared with a fruit without a compet-
ing regrowth. Fruit DM was reduced from 17.3
to 14.9, a 14% reduction. Regrowth FW was
reduced by 24%, while regrowth DW reduced
by 23%. Hence, competition had a larger
relative effect on fruit DW than on fruit FW,
and the vegetative sink similarly suffered effects
on both FW and DW. The relative reduction in
FW of the fruit and regrowth were similar but
the fruit suffered greater relative DW loss than
regrowth.
In year 2, the fruit regrowth competition
experiment was repeated, with the addition of
two fruits in competition (Table 3). Compar-
ison between the same treatments in year 1 and
year 2, which involved different orchard blocks,
showed no significant difference (pB0.05) in
fruit and vegetative FW and DW results at final
harvest, ie, about 180 DAFB (Table 2 and 3).
With two fruits, (year 2) at 181 DAMB each
of the fruits had similar FW and DW. Compe-
tition between two fruits caused growth of each
fruit to be reduced by 15% in FW and 23% in
DW, resulting in 12% loss of DM. Competition
of a single fruit with regrowth in year 2 resulted
in a loss of fruit FW (28%) and DW (39%)
resulting in a loss of 15% DM. Looked at in a
different way, two fruits resulted in 268.0 g FW
and 41.6 g DW while one fruit plus regrowth at
harvest had a FW of 183.3 g and DW of 39.4 g.
The total FW of two fruits was greater than
that of one fruit plus a deleafed regrowth,
but the DWs were not significantly different
(p B0.05).
Discussion
Lai et al. (1989) and Hopping (1990) reported
that fruit weight of kiwifruit increases with the
number of seeds in the fruit. In our experi-
ments, the fruit were well pollinated, as fruit
averaged more than 1000 seeds per fruit,
considered adequate for commercial produc-
tion. Seed counts per fruit taken at harvest did
not differ significantly among the treatments
(Table 1), indicating that the observed differ-
ences in fruit FW and DM were not caused by
differences in seed counts. Previous studies
have shown that seed number has much less
influence on fruit size when fruit are supplied
with adequate resources on girdled shoots
(Snelgar & Thorp, 1988). Lai et al. (1988)

Table 3 Year 2, data at 180 DAMB. Average fruit and regrowth fresh weight (FW), dry weight (DW) and dry
matter content (DM), with standard errors (se) in parenthesis, for treatments with four source leaves (4L),
one or two fruits (1F, 2F), and regrowth (R) in year 2. Means differ significantly (P B0.05) when the
difference is more than 1.5 times the sum of the standard errors (Payton et al. 2003)

Fruit Regrowth

Treatment FW (g) DW (g) DM (%) FW (g) DW (g) DM (%)

4L1F 157.8 (6.7) 27.1 (1.6) 17.2 (0.5)

4L2F 133.1 (4.6) 20.6 (1.4) 15.1 (0.6)
134.9 (6.2) 21.0 (1.6) 15.1 (0.7)
4L1FR 113.0 (5.7) 16.5 (1.3) 14.6 (0.7) 70.3 (4.6) 22.9 (2.2) 31.7 (1.5)
108 PEH Minchin et al.
showed that phloem transport within a shoot of
kiwifruit follows an orthostichy relation of the
nth node supplying nodes n5 and n8, so we
may expect this orthostichy relationship to
affect the distribution of photosynthate within
our treatments. However, Lai et al. found that
on pruning, this relation was completely lost.
Hence, because our shoot was pruned, issues of
orthostichy are not expected to be relevant. We
saw a consistent increase in fruit size with the
number of source leaves (Table 1).
Source limitations
At harvest, one fruit supplied by seven leaves
resulted in fruit with larger FW and DW than
when supplied by four leaves (Table 1), con-
firming that four leaves were not sufficient to
saturate fruit growth. Reducing the number of
source leaves tended to increase the variability
in FW and DW, seen by the increasing
standard error of these measurements. This
was not unexpected. With a specific number
of leaves, source supply would be expected to
vary between replicates due to differences in
total leaf area and in light exposure. With a
large number of leaves on a shoot, carbohy-
drate supply would approach saturation of sink
utilization. Hence, variation in source supply
between replicates of a treatment with a small
number of leaves is expected to result in greater
variability on FW and DW than between
replicates with a larger number of leaves.
The effect of leaf number on fruit and
regrowth FW and DW is shown in Fig. 3.
The rate of increase of both fruit FW and DW
fell with increasing leaf number, suggesting
saturation as carbohydrate supply increased.
Regrowth did not show signs of saturation,
increasing linearly with leaf number. Since fruit
FW and DW increased with leaf number at
different rates, fruit DM must change with the
number of supply leaves. Put another way, on
increasing source supply (i.e. leaf number) a
bigger fraction of the increased supply went
into DM growth of the vegetative sink than
went into DM growth of the fruit. The DM
of a fruit increased with increasing number
of leaves. The major component of FW is
water, imported into a sink by phloem and
xylem flow, but DW is predominantly imported
by the phloem. These two flow systems are
not independent. Their interaction is complex,
depending on transpirational water loss
through the fruit surface and osmotic effects
within the fruit. The osmotic effects are depen-
dent on photosynthate import, its route of
phloem unloading and metabolic fate. Interac-
tions between phloem and xylem flow are only
just starting to be investigated (see Holbrook &
Zwieniecki 2005) and to date this has probably
best been done in the modelling of peach fruit
growth (Fishman & Génard 1998; Génard et al.
1998; Lescourret et al. 1998; Lescourret &
Génard 2005). This level of mechanistic detail
has not been developed for other fruit.
Standard commercial orchard practice
produces fruit of about 95
115 g FW and
about 14
17 DM which corresponds to two to
three leaves per fruit (see Fig. 3) with a girdled
shoot. Lai et al. (1989) found that two leaves
per fruit on a girdled shoot resulted in similar
fruit size to fruit on non-girdled shoots, so this
is consistent with our observations. Extremely
large fruit can be produced by higher leaf
numbers. With seven leaves we achieved a fruit
FW of 174 g with 18.4 DM, demonstrating the
potential of fruit size.
Fruit growth curves have often been inter-
preted in terms of three phases of growth
(Coombe 1976), which are a direct consequence
of double sigmoid growth, which has often
been reported (Coombe 1976) and first
reported for kiwifruit by Hopping (1976). The
three growth stages are: stage 1 and 3 when
each growth period is at its peak and stage 2
being the time between the two growth periods
when the cumulative growth appears to slow
and then increase again. To a variable degree,
the time courses of fruit FW growth showed
the three stages of fruit growth. As pointed out
by Coombe (1976), description of fruit growth
by means of sigmoid curves is problematic as
the observed shape depends strongly on the
method, and accuracy, of measurement as
well as the delay between the individual times
of more rapid growth which can obscure or
emphasize phases 2 and 3. Coombe (1976)
suggested that a more rigorous description
based on curve fitting should be used, but this
has not been done in the fruit literature.
Numerical differentiation of the observed
 Competition between fruit and vegetative growth in Hayward kiwifruit 109
growth curve can be revealing, but great care is
needed with numerical differentiation as this
amplifies the noise on what are already typi-
cally noisy observations. During anthesis there
is some cell division and this rapidly increases
immediately after anthesis, lasting for a limited
period, and is associated with stage 1 of fruit
growth (Coombe 1976). Then cell enlargement
occurs which is traditionally associated with
stage 3 of growth. The extent of stage 2
depends on the delay between the two stages
of rapid growth. In our data, the two rapid
growth stages were obvious only with four
source leaves. With four leaves, growth rate
showed a clear stage 2 and this was accentuated
when the fruit were competing with the vege-
tative sink, and much less marked when the
fruit was supplied by seven leaves. With two
leaves, stage 1 growth was much reduced and
there was no sign of stage 3 growth, with FW
growth constant from about 90 DAMB up to
harvest at 181 DAMB. The two-leaf treatment
had higher growth rate at 180 DAMB than any
of the other treatments.
The time course of regrowth extension is
shown in Fig. 2B, with regrowth increasing
with carbohydrate supply. At 181 DAMB,
regrowth supported by seven leaves was about
52% greater than when supported by four
leaves. Vegetative regrowth continued until
about 100 DAMB, after which there was little
extension growth. Elongation was reducing at
the time of onset of phase 3 fruit growth.
Hence, phase 3 of fruit growth may be a direct
consequence of increasing availability of carbo-
hydrate through reducing vegetative demand.
The pattern of regrowth extension was
quite different from that of its DW and is
quite misleading as to its time course of
carbohydrate utilization. This is readily seen
in Table 2. Here we see that between 100 and
181 DAMB, when in competition with a single
fruit, regrowth FW did not significantly in-
crease but there was a marked increase in its
DW. Hence, although there was no length
increase, there was a considerable increase in
regrowth DW. This could not be accounted for
by increase in regrowth diameter, as the time
course of regrowth volume was the same as that
of length (data not presented). By 100 DAMB,
fruit DW growth had been considerably
impaired by the competition from the regrowth
(a loss of 8.5 g) and by 181 DAMB the loss
was only slightly larger (9.6 g).
Sink competition
In the first 100 DAMB, vegetative growth was
reduced less by the presence of a fruit than fruit
growth was reduced by the presence of a
regrowth (Table 2). That is, vegetative growth
had a greater ability to attract growth sub-
strates than did the fruit. When there were two
fruits competing for available resources, the
two fruits at 180 DAMB were indistinguishable
(Table 3). The total DW growth resulting with
two fruits was the same as one fruit plus a
regrowth. Thus, DW growth was unaffected by
the sink type even though the FW distribution
between sink types was not equal. The vegeta-
tive sink had a higher priority for carbohydrate
supply than did the fruit. While this describes
the data, it does not explain the result.
Competition by a fruit sink is different from
competition by a vegetative sink.
Currently, the only mechanistic explanation
of differences in sink types is by differences
in the Michaelis-Menton description of the
saturatable kinetics of sink utilization of
available carbohydrate (Minchin et al. 1993;
Lacointe & Minchin 2008). In this description,
sink function was described by saturatable
Michaelis-Menton kinetics and differences in
priority result for differences in Vmax. When the
km of both sinks was assumed to be the same,
the sink with the lower Vmax had the higher
priority. If various sinks utilize the same carrier,
or enzymology, then the effective km describing
this will be the same as the Michaelis-Menton
parameter km that describes carrier binding.
For different sink types, the effective kms are
expected to be different, so differences in Vmax
may not be the sole cause of differences in sink
priority between fruit and vegetative sinks.
Equivalent sinks, that is sinks of the same
physiological type, described by the same km
and their individual Vm’s, scaled according to
their individual flow resistances resulted in this
model having the same priorities (Minchin et al.
1993) i.e., with a change in supply the fraction of
supply transported into each sink remained
fixed. This is consistent with equal growth seen
110 PEH Minchin et al.
in the two fruits, but not between fruit and
regrowth which would be modelled as non-
equivalent sinks.
From about 100 DAMB, FW growth of the
vegetative sink was small, practically zero in the
presence of a competing fruit, and increasing
by 15% without this competition (Table 2).
But, both fruit and regrowth DW continued to
increase, with the fruit gaining almost twice the
DW increment of the regrowth. Consequently,
the fruit DM showed a large increase between
100 and 181 DAMB.
Field interpretation
The defoliated regrowth used in this work is
quite unrepresentative of normal vegetative
growth, but was used here as a means of
maintaining the vegetative growth as a sink.
Normal vegetative growth starts as a net
carbohydrate sink and as leaves mature
becomes a net exporter and supplies the fruit:
a series of experiments to explore the dynamics
of this more complex system are currently being
carried out.
This work investigated the competition
between carbohydrate sinks in kiwifruit vines.
We were interested in the plant responses at the
organ level, and possible vine management
issues. Girdling, as a means of inhibiting
phloem transport, has been used as a horticul-
tural practice for thousands of years (Goren
et al. 2004) and is still not well understood.
It appears to induce a complex interaction
between changes in endogenous hormones
and carbohydrate distribution (Goren et al.
2004; Theron & Steyn 2008). At this stage, we
have not attempted to give a complete physio-
logical interpretation to the reported responses.
This will be done in the near future through
application of a functional-structural-plant
model of kiwifruit, incorporating phloem and
xylem transport to model proposed mecha-
nisms of sink competition (Lacointe & Minchin
2008; Cieslak et al. 2009).
Conclusions
The aim of this work was to investigate the
interactions between fruit and between fruit
and vegetative growth in ‘Hayward’ kiwifruit,
to increase understanding of sink competition
and how this can be used to optimize fruit FW
and DM.
We have shown that fruit FW and DW at
harvest are readily varied by carbohydrate
supply level, but not equally. As supply in-
creased, DW increases proportionally faster
than FW, resulting in increased DM. We have
shown that competition for available carbohy-
drate by a regrowth has a large detrimental
impact on both fruit FW and DM. Pruning
that stimulates regrowth can be expected to
have a large negative impact on the fruit
properties at harvest. A regrowth has a much
bigger effect than an extra fruit. The total DW
imported by two fruits or one fruit plus the
defoliated regrowth were equal but a fruit had
greater FW growth than that of a defoliated
regrowth.
These quantitative data will provide impor-
tant calibration data for both mechanistic
models of source sink interactions as well as
of fruit growth.
Acknowledgements
This work was funded by Plant & Food Research’s
internal kiwifruit reinvestment fund and FRST
contract C06X0706.
References
Austin PT, Hall AJ, Gandar PW, Warrington IJ,
Fulton TA, Halligan EA 1999. A compartmental
model of the effect of early-season temperature
on potential size and growth of ‘Delicious’ apple
fruits. Annals of Botany 83: 129
143.
/
Avery DJ, Priestley CA, Treharne KJ 1979. Integration
of assimilation and carbohydrate utilization in
apple. In: Marcelle R, et al eds. Photosynthesis
and plant development. The Hague, Junk. Pp.
221
231.
BioGro 2009. BioBro Manual, BioGro Society, PO
Box 9693, Marion Square, Wellington, NZ.
http://www.biogro.co.nz/main.php?page230
Burdon J, McLeod D, Lallu N, Gamble J, Petley M,
Gunson A 2004. Consumer evaluation of ‘Hay-
ward’ kiwifruit of different at-harvest dry matter
contents. Postharvest Biology and Technology /
34: 245
255.
 Competition between fruit and vegetative growth in Hayward kiwifruit 111
Cieslak M, Seleznyova AN, Hanan J 2009. A
functional-structural kiwifruit vine model.
Proceedings of the Third International Sympo-
sium on Plant Growth Modelling, Simulation,
Visualization and Applications. Beijing (China),
9
13 November 2009.
Coombe BG 1976. The development of fleshy fruits.
Annual Review of Plant Physiology 27: 507
/
528.
Crisosto CH, Crisosto GM 2001. Understanding
consumer acceptance of early harvested ‘Hayward’
kiwifruit. Postharvest Biology and Technology
22: 205
213.
Fishman S, Génard M 1998. A biophysical model of
fruit growth: simulation of seasonal and diurnal
dynamics of mass. Plant. Cell and Environment
21: 739
752.
Génard M, Lescourret F, Ben Mimoun M, Besset J,
Bussi C 1998. A simulation model of growth
at the shoot-bearing fruit level. II. Test and
effect of source and sink factors in the case
of peach. European Journal of Agronomy 9:
189
202.
Goren F, Huberman M, Goldschmidt EE 2004.
Girdling: physiological and horticultural aspects.
Horticultural Reviews 30: 1
38.
/
Greer DH 1999. Seasonal and daily changes in
carbon acquisition of kiwifruit leaves with and
without auxiliary fruit. New Zealand Journal of
Crop and Horticultural Science 27: 23
31.
/
Greer DH, Jeffares D 1998. Temperature-dependence
of carbon acquisition and demand in relation to
shoot growth of kiwifruit (Actinidia deliciosa)
vines grown in controlled environments. Aus-
tralian Journal of Plant Physiology 25: 843
850.
/
Grossman YL, DeJong TM 1994. PEACH: A
simulation model of reproductive and vegetative
growth in peach trees. Tree Physiology 14: /
329
345.
Grossman YL, DeJong TM 1995. Maximum fruit
growth potential and seasonal patterns of
resource dynamics during peach growth. Annals
of Botany 75: 553
560.
/
Harker FR, Carr BT, Lenjo M, MacRae EA,
Wismer WV, Marsh KB, Williams M, White
A, Lund CM, Walker SB, Gunson FA, Pereira
RB 2009. Consumer liking for kiwifruit flavour:
A meta-analysis of five studies on fruit quality.
Food Quality and Preference 20: 30
41.
/
Holbrook NM, Zwieniecki MA 2005. Vascular
transport in plants. New York, Elsevier
Academic Press.
Hopping ME 1976. Structure and development of
fruit and seeds in Chinese gooseberry (Actinidia
chinensis Planch.). New Zealand Journal of
Botany 14: 63
68.
/
Hopping ME 1990. Floral biology, pollination, and
fruit set. In: Warrington IJ, Weston GC eds.
Kiwifruit: science and management. Auckland,
NZ, Ray Richards. Pp. 71
96.
Jordan RB, Walton EF, Klages KU, Seelye RJ 2000.
Postharvest fruit density as an indicator of dry
matter and ripened soluble solids of kiwifruit.
Postharvest Biology and Technology 20: 163
/
173.
/
Lacointe A, Minchin PEH 2008. Modelling phloem
and xylem transport within a complex architec-
ture. Functional Plant Biology 35: 772
780.
/
Lai R, Wooley DJ, Lawes GS 1988. Patterns of
/
assimilate transport from leaves to fruit within a
kiwifruit (Actinidia deliciosa) lateral. Journal of
Horticultural Science 63: 725
730.
/
Lai R, Wooley DJ, Lawes GS 1989. Effect of leaf to
fruit ratio on fruit growth of kiwifruit (Actinidia
deliciosa). Scientia Horticulturae 39: 247
255.
/
/
Lancaster J 2002. What makes a good flavoured
kiwifruit? New Zealand Kiwifruit 149: 10
11. /
Lescourret F, Ben Mimoun M, Génard M 1998. A
simulation model of growth at the shoot-
bearing-fruit level. I Description and parame-
terisation for peach. European Journal of
Agronomy 9: 173
188.
/
Lescourret F, Génard M 2005. A virtual peach fruit
model simulating changes in fruit quality during
the final stage of fruit growth. Tree Physiology
25: 1303
1315.
/
Minchin PEH, Thorpe MR, Farrar JF 1993. A
simple mechanistic model of phloem transport
which explains sink priority. Journal of Experi-
mental Botany 44: 947
955.
/
Minchin PEH, Richardson AC, Patterson KJ,
Martin PJ 2003. Prediction of final weight
for Actinidia chinensis ‘Hort16A’ fruit. New
Zealand Journal of Crop and Horticultural
Science 31: 147
157.
/
Palmer JW 1992. Effects of varying crop-load on
photosynthesis, dry matter production and
partitioning of Crispin/M.27 apple trees. Tree
Physiology 11: 19
33./
Payton ME, Greenstone MH, Schenker N 2003.
Overlapping confidence intervals or standard
error intervals: What do they mean in terms of
statistical significance? Journal of Insect Science
3:34 (available online: insectscience.org/3:34).
Quilot B, Génard M, Lescourret F, Kervelle L 2005.
Simulating genotypic variation of fruit quality
in an advanced peach x Purus davidiana cross.
Journal of Experimental Botany 56: 3071
3081.
/
112 PEH Minchin et al.
Richardson AC, McAneney KJ, Dawson TE 1997.
Carbohydrate dynamics in kiwifruit. Journal of
Horticultural Science 72: 907
917.
/
Snelgar WP, Hall AJ, Ferguson AR, Blattmann P
2005. Temperature influences growth and
maturation of fruit on ‘Hayward’ kiwifruit
vines. Functional Plant Biology 32: 1
12.
/
Snelgar WP, Thorp TG 1988. Leaf area, final fruit
weight and productivity in kiwifruit. Scientia
Horticulturae 36: 241
249.
Snelgar WP, Thorp TG, Patterson KJ 1986. Optimal
leaf: fruit ratios for fruit growth in kiwifruit.
Acta Horticulturae 175: 115
120.
/
Snelgar WP, Manson PJ, Martin PJ 1992. Influence
of time of shading on flowering and yield of
kiwifruit vines. Journal of Horticultural Science
67: 481
487.
/
Taylor JA, Praat JP, Bollen AF 2007. Spatial
variability of kiwifruit quality in orchards and
its implications for sampling and mapping.
HortScience 42: 246
250.
/
Theron KI, Steyn WJ 2008. Girdling: science behind
the age-old technique. Acta Horticulturae 800:
/
51
60.
Waring PF, Patrick J 1975. Source-sink relations
and the partition of assimilates in the plant. In:
Cooper JP ed. Photosynthesis and productivity
in different environments. Cambridge, UK,
Cambridge University Press.
Wardlaw IF 1990. The control of carbon partition-
ing in plants. New Phytologist 116: 341
81.
/
Woodward TJ, Green TGA, Clearwater MJ 2007.
Between-vine variation in ‘Hayward’ kiwifruit
vine income: The use of Lorenz curves and
Gini coefficients to describe variation and
assess the potential for zonal management.
European Journal of Horticultural Science 72: /
280
287.
Wright CJ 1989. Interactions between vegetative and
reproductive growth. In: Wright CJ ed. Manip-
ulation of fruiting. London, UK, Butterworth.