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New Zealand Journal of Crop and Horticultural Science

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

Physiological responses of kiwifruit vines (Actinidia

chinensis Planch. var. chinensis) to trunk girdling
and root pruning

MZ Black , KJ Patterson , KS Gould & MJ Clearwater

To cite this article: MZ Black , KJ Patterson , KS Gould & MJ Clearwater (2012) Physiological
responses of kiwifruit vines (Actinidia�chinensis Planch. var. chinensis) to trunk girdling and
root pruning, New Zealand Journal of Crop and Horticultural Science, 40:1, 31-41, DOI:

To link to this article: https://doi.org/10.1080/01140671.2011.603343

Published online: 26 Sep 2011.

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New Zealand Journal of Crop and Horticultural Science
Vol. 40, No. 1, March 2012, 3141

Physiological responses of kiwifruit vines (Actinidia chinensis Planch. var.

chinensis) to trunk girdling and root pruning
MZ Blacka*, KJ Pattersonb, KS Gouldc and MJ Clearwaterd
The New Zealand Institute for Plant & Food Research Ltd (Plant & Food Research), Te Puke Research
Centre, Te Puke, New Zealand; bPlant & Food Research, Mt Albert Research Centre, Auckland, New Zealand;
School of Biological Sciences, Victoria University of Wellington, New Zealand; dSchool of Biological Sciences,
University of Waikato, Hamilton, New Zealand
(Received 3 May 2011; final version received 22 June 2011)

Hydraulic conductance and photosynthesis were measured in kiwifruit (Actinidia sp.) vines to
observe responses to the horticultural practices of trunk girdling and root pruning. Little is
understood regarding the physiological effects of these practices in kiwifruit. Combined
reductions in hydraulic conductance and photosynthesis were expected following root pruning.
The primary response to trunk girdling was expected to be photosynthetic. Measurements of
transpiration and xylem pressure potentials (C) were used to calculate hydraulic conductance
(K). Stomatal conductance (gs) and photosynthesis (A) were measured periodically. Neither
treatment affected fruit growth. The response to root pruning was primarily hydraulic with
significant reductions in midday C indicating reduced K; reductions in A and gs were small.
Girdling resulted in significant reductions in gs and A, with little effect on C or K. Reduced
A following girdling was considered to be a consequence of stomatal and biochemical limitation,
with biochemical down-regulation following reduced carbohydrate demand.
Keywords: Actinidia chinensis; hydraulic conductance; kiwifruit; photosynthesis; root pruning;
trunk girdling

Introduction 5 mm in width, from around the circumference

In order to meet quality standards set by of the trunk, avoiding damage to the xylem.
kiwifruit (Actinidia sp.) marketers, New Trunk girdling temporarily increases the
Zealand kiwifruit growers have adopted a range availability of carbohydrates to the developing
of innovative orchard practices. Two such fruit through disruption of the source:sink ratio
practices, trunk girdling and root pruning, are of the vine, by temporarily removing the root
quite invasive to the plant and raise questions system as a carbohydrate sink (Noel 1970).
regarding how plant function is affected. Trunk The technique was adopted by New Zealand
girdling and root pruning were introduced to kiwifruit growers in the early 2000s, following
New Zealand kiwifruit orchards with the aim of demonstration of its success in a number of
increasing fruit size and percentage dry matter other fruit crops (Goren et al. 2003). In grapes,
(DM), with DM being associated with fruit for example, berries are larger and mature
sweetness (Harker et al. 2009). earlier (Weaver 1976; Williams et al. 2000).
Trunk girdling involves the removal of a Consistent positive effects on fruit size and DM
band of bark and phloem tissue, approximately content have been achieved as a consequence of

*Corresponding author. Email: mary.black@plantandfood.co.nz

ISSN 0114-0671 print/ISSN 1175-8783 online
# 2012 The Royal Society of New Zealand
32 MZ Black et al.

trunk girdling kiwifruit vines at particular a photosynthetic rather than hydraulic

times of the year (Currie et al. 2005) and the response. In oak, however, trunk girdling
technique is now widely used. reduced xylem sap flux within 2 days of treat-
Root pruning entails a tractor-drawn blade, ment (De Schepper et al. 2010). The observed
approximately 300400 mm in length (Schupp reduction was too rapid to have resulted from
& Ferree 1987; Kurtural & Taylor 2004), being effects of limited carbohydrate supply on root
dragged through the soil, along the row, within system size. Reduced water use following trunk
450600 mm of the trunk (Ferree et al. 2000; girdling has been attributed to an indirect
Kurtural & Taylor 2004). It may be suggested response, with reduced transpiration related to
that root pruning has a temporary effect on stomatal limitation of photosynthesis (Williams
carbohydrate partitioning, with the reduced & Ayars 2005; De Schepper et al. 2010).
size of root system requiring a smaller fraction Root pruning has an immediate, long-
of carbohydrates (Detling et al. 1980). Root term impact on the size of the root system,
pruning is a more recent introduction to the the severity of which is determined by the
kiwifruit industry; trials have shown consistent proportion of root system removed. Reducing
increases in fruit DM content and the potential root surface area will disrupt the plant’s
to increase fruit size (Patterson et al. 2009), carbohydrate balance, as well as affecting water
promoting interest in the technique. In addi- uptake capacity. The effects of root pruning on
tion, root pruning has the potential to reduce plant hydraulics and carbon assimilation are
vegetative vigour of the kiwifruit vine, as has not clearly understood. The reduction in root
been observed in other fruit crops (Ferree 1990; area is thought to reduce root system demand
Maas 2008). for carbohydrates, making more available to
Previous research into the use of these other organs. Consistent with that hypothesis
techniques in kiwifruit has studied the effects are observations of larger fruit and increases in
on fruit quality and growth (Currie et al. 2008; fruit carbohydrate content (Patterson et al.
Patterson et al. 2009); however, little is known 2009). In contrast, vegetative growth is often
regarding the effects on physiological processes reduced; this reduction in vegetative growth
within kiwifruit vines. Trunk girdling and root is unlikely to occur through carbohydrate
pruning both have an impact on the root system. limitation; therefore, it may be suggested that
Following trunk girdling, reductions in root a hydraulic effect is limiting vegetative growth.
carbohydrate reserves have been observed in Reductions in stomatal conductance, transpira-
peach and kiwifruit (Jordan & Habib 1996; tion and leaf water potentials have been found
Currie et al. 2005), along with reductions in in apple, grape, pear and peach (Geisler &
fine root production in Japanese Persimmon Ferree 1984; Poni et al. 1992; Smart et al. 2006),
(Fumuro 1997). Annual girdling and the confirming a hydraulic response following root
application of multiple girdles in a single season pruning.
may magnify these effects in the long term. The The objective of this experiment was to
nature of the physiological response to trunk compare the physiological responses of kiwi-
girdling is not clear. Correctly applied girdles fruit vines to trunk girdling and root pruning.
should not cause substantial damage to the We hypothesized that in kiwifruit, trunk
xylem and should therefore not directly affect girdling manipulates carbohydrate partitioning,
hydraulic conductance. In grape, reductions thereby reducing sink demand and resulting
in stomatal conductance have been observed in reduced photosynthesis. Photosynthetic
following trunk girdling (Roper & Williams reductions could involve both biochemical and
1989; Williams et al. 2000), although this was stomatal limitations. We further hypothesized
not associated with reductions in leaf water that root pruning would moderate photosynth-
potential (Williams et al. 2000), which suggests esis through reduced stomatal conductance. It
Vine responses to girdling and root pruning 33

was postulated that a reduction in root area equally distributed throughout the day. The
would result in rapid and significant reductions pots were covered with transparent plastic to
in hydraulic conductance, stomatal conduc- prevent soil evaporation.
tance and transpiration. Treatments were applied on 12 February
2009, with three replicates, ensuring vines from
the same treatment were not positioned next to
Materials and methods
each other. Timing of treatment application
Plant material was chosen to coincide with one of the recom-
An experiment was conducted in a plastic mended commercial timings for trunk girdling
greenhouse at the Plant & Food Research Te (Currie et al. 2008). Trunk girdling involved the
Puke research orchard, New Zealand. Vines use of a modified blunt chainsaw blade to
were 9 year-old yellow kiwifruit (Actinidia remove a band of bark including the phloem
chinensis Planch. var. chinensis ‘Hort16A’) tissue, approximately 5 mm in width, from
grafted on to open pollinated green kiwifruit around the trunk. Care was taken to ensure
(Actinidia deliciosa (A.Chev.) C.F. Liang et that all phloem tissue was removed while
A.R. Ferguson var. deliciosa) rootstocks. minimizing damage to the xylem tissue. Root
Grafted seedlings were transferred to 100 L pruning was performed by cutting vertically
fabric root control bags (Smart Pot*In to the base of the pot, along two sides of
GroundTM), before being planted out in the the trunk, approximately 150 mm away from
orchard. The root control bags were buried the trunk, with an electric reciprocating
and vines were maintained for approximately saw (CR18DS, Hitachi, Tokyo, Japan). This
7 years, managed according to standard removed approximately 50% of the root ball;
practice for the cultivar, adapted from re- the severed portion was left in place.
commendations for A. deliciosa (Sale & Lyford
1990). The annual fertilizer regime for the vines
included a base application of potassium, Hydraulic conductance
phosphate, sulphur and magnesium in October Transpiration (Eplant) was estimated based
and applications of nitrogen in November, on weight loss after correcting for drainage.
January and July. Vines were irrigated with Drainage from each pot was measured using
individual sprinklers in close proximity to individual tipping bucket rain gauges (Rain-O-
the trunk. The vines were trained to a T-bar Matic, Scott Technical Instruments, Hamilton,
support system. New Zealand). Weighing lysimeters consisted
of a load-cell (Model 60060C1000KG-300M,
Sensortronics, California, US) sandwiched
Experimental conditions between two 1 m2 aluminium plates; load-cells
In November 2008, approximately 1 month and rain gauges were connected to a datalogger
post-anthesis, plants were transferred from the via a multiplexer and a switch closure input
ground, remaining in the root control bags, to module, respectively (CR10X; AM416; SDM
custom-made 150 L pots. Pots were fitted with SW8A, Campbell Scientific, Utah, US). Average
a drainage tube and a metal framework for weight of each potted vine and total drainage
supporting the trunk and leader. Vines were were recorded at 15 min intervals.
left to acclimatize outdoors for 3 weeks, Xylem pressure potentials (C, MPa) were
before being transferred to individual weighing measured using a pressure chamber (model
lysimeters in the plastic greenhouse. Fans were 3005, Soil Moisture Equipment Corp, Santa
used to assist natural air flow ventilation. Pots Barbara, US); measurements were made on
were irrigated with individual sprinklers; vines three leaves per vine at pre-dawn, on uncovered
received three applications of 6 L of water, leaves (C soil) and at midday on both covered
34 MZ Black et al.

(C shoot) and uncovered (C leaf) leaves. Pre-dawn expanded, sunlit leaves per vine, using natural
measurements were assumed to be equivalent to light.
soil water potential (C soil) (Ritchie & Hinckley Chamber CO2 concentration was controlled at
1975). Midday assessments of non-transpiring 400 mmol mol 1 and vapour pressure deficit was
covered leaves approximate xylem pressure maintained at an average of 2.5 kPa. Leaf
within the shoot (Meinzer et al. 2001). Covered samples were collected for d13C isotope analysis
leaves were enclosed in foil coated plastic bags, from each vine at weekly intervals; two samples
a minimum of 2 hours prior to measurement. were taken per vine. Each sample comprised six
Midday C shoot and C leaf were measured on newly emerging leaves selected from each vine,
clear sunny days, between 11:00am and based on the assumption that expanding leaves
3:00pm, using mature fully expanded leaves; are representative of recently fixed carbon
measurements were made at six timings post- (Meinzer et al. 1993). Samples were dried at
treatment. Soil water potential was assessed at 80 8C to a constant weight; finely ground and
four timings post-treatment. 2.8 mg samples weighed into foil cups. A mass
Hydraulic conductance was calculated for spectrometer (20/20 isotope analyser, Europa
the soil to shoot (Kshoot) and soil to leaf (Kplant) Scientific, Crewe, UK) at the University of
pathways as: Waikato Stable Isotope Unit was used to
measure d13C () discrimination in comparison
K ¼ Eplant =DW (1) with the Pee Dee belemnite standard.
where Eplant is the mean rate of Eplant between
9:00am and 3:00pm per vine for the days when Fruit measurements
C was measured. ^C is the difference between
Percentage fruit DM was assessed at weekly
C at points along the hydraulic pathway (C soil
intervals, from samples of four fruit per vine.
 C shoot for Kshoot and C soil  C leaf for Kplant).
Actual fruit fresh weight (FW), fruit length
Sapwood area specific conductance (Ks) was
and maximum and minimum diameters at
calculated to account for differences in trunk
the equatorial region were recorded for each
cross-sectional area:
fruit. A 2 mm-thick cross-sectional slice was
Ks ¼ K=Asw (2) taken from the equatorial region of each fruit,
weighed and dried in a dehydrator at 65 8C
Trunk sapwood area was estimated from until constant weight was achieved. Dried slices
measurements of trunk diameter inclusive of were reweighed (DW) and DM content was
bark and bark thickness at points around the calculated as:
trunk, above and below the visible graft union.
Sapwood area (Asw, cm2) for rootstock and
scion was calculated, assuming all tissue below DM ¼ ðDW =FW Þ  100 (3)
the bark was conducting.
Length and diameter measurements (mm) and
actual FW (g) of the sampled fruit were used in a
Photosynthesis and stomatal conductance regression analysis to obtain a calibrated multi-
Stomatal conductance (gs) and net photo- plier for non-destructive fruit FW estimates as
synthetic rate (A) were measured and leaf inter- described by Minchin et al. (2003). The slope of
nal CO2 content (ci) calculated using a portable the regression line, used as the multiplier was
gas exchange system (LI-6400, LI-COR, 0.54, with an R2 of 97.5%. Fruit length (L)
Nebraska, US), at times to coincide with and maximum and minimum diameters at
measurements of Cshoot and C leaf. Measure- the equatorial region (Dmax and Dmin, respec-
ments were taken on four to six mature, fully tively) were measured weekly post-treatment on
Vine responses to girdling and root pruning 35

six marked fruit per vine and fresh weight and C leaf (9SEM) in root-pruned vines were
calculated as: approximately 0.290.02 MPa lower than in
control vines.
FW ¼ ðððL  Dmax  Dmin Þ=1000Þ  0:54Þ (4) Midday transpiration in control vines
averaged 2.0 kg s 1 104, reaching a maximum
rate of 3.2 kg s 1 104. All vines demonstrated
Statistical analysis a decline in Eplant 1 day prior to treatment, which
Statistical analyses were carried out in R version was maintained throughout the experiment.
2.10.01 (Comprehensive R Archive Network, A treatment effect on Eplant was not observed
cran.r-project.org). Data were tested for normal (Fig. 1A). In general, Kshoot and Kplant (Table 2)
distribution using the Shapiro-Wilk test and in trunk-girdled vines remained comparable to
log transformations performed to normalize controls. Maintenance of Eplant, combined with
data where required for ANOVA analyses. an increase in the soil to shoot pressure gradient
Treatment differences were tested using in root-pruned vines, indicated reductions in
ANOVA. Pairwise Least Significant Difference Kshoot (Fig. 1B). Reductions in Kshoot (P B0.05)
(LSD) tests were used to compare treatments were evident during the 2 weeks post-treatment;
where significant differences were found. however, treatment effects were not maintained
Dieback of a single root-pruned vine towards to 4 and 5 weeks post-treatment. Whole
the end of the study led to the exclusion of data vine hydraulic conductance was less affected
from this vine. (Table 2), with reductions, although insignif-
icant, only evident for the first week post-
treatment. Sapwood area was similar for all
Results vines, averaging 14.090.6 cm2 for rootstock
Effects on C differed with treatment (Table 1). and 12.490.6 cm2 for scion cross-sections.
Pre-dawn xylem pressure potential was between Whole vine sapwood area specific conductance
0.01 to 0.1 MPa, remaining comparable in (Ks-plant) showed close agreement with Kplant,
control and trunk-girdled vines. Reductions with no significant differences between
were evident in root-pruned vines. Midday treatments. Significant reductions in soil to
Cshoot and Cleaf in trunk-girdled vines were shoot sapwood area specific conductance (Ks-
comparable to controls. In root-pruned vines, shoot) were observed in root-pruned vines, show-
midday C shoot and C leaf were significantly ing agreement with Kshoot (Table 2).
reduced within 1 day of treatment, with reduc- Trunk girdling had a larger effect on A
tions maintained for 7 weeks. Midday C shoot (Fig. 2A) and gs, (Fig. 2B) than root pruning,

Table 1 Post-treatment xylem pressure potentials (MPa) for pre-dawn leaf (Csoil), midday kiwifruit shoot
(Cshoot) and leaf (C leaf). Data for Csoil are means n 3 vines at three timings (n 2 vines for root pruned),
data for C shoot and C leaf are means n 3 vines at seven timings (n 2 vines for root-pruned), 91SEM.
Values in the same column sharing the same letter are not significantly different (P B0.05). Xylem pressure
potentials were measured with the pressure chamber at pre-dawn (C soil) and at midday on covered non-
transpiring leaves (Cshoot) and on openly transpiring leaves (Cleaf).

Csoil C shoot C leaf


Control 0.0490.003 a 0.3490.01 a 0.5490.02 a

Root pruned 0.0590.005 b 0.5290.02 b 0.7190.02 b
Trunk girdled 0.0390.003 a 0.3690.02 a 0.5990.03 a
P value 0.0001 B0.01 B0.01
36 MZ Black et al.

with control vines, although not significantly.

Final measurements, at 5 and 7 weeks post-
treatment, showed an increase to rates higher
than that of control vines. Leaf internal CO2
content of root-pruned vines was maintained
at levels comparable to controls. Trunk-girdled
vines demonstrated consistently higher d13C
than control vines, with significant differences
evident at 4 weeks post-treatment (Fig. 3).
Leaf sample analysis of d13C content from 1
to 5 weeks post-treatment showed little differ-
ence between control and root-pruned vines.
Mean fruit weight increased from 72.19
2.0 g to 85.391.8 g over the 7 week period,
with no significant differences between treat-
ments (Fig. 4). Rate of fruit growth (g day 1),
calculated from FW estimates using equation
(4), showed a steady decline for all vines
throughout the experiment, with little evidence
Figure 1 A, Average transpiration rate (kg s 1 104)
of a treatment response (data not shown).
between 9:00am and 3:00pm for root-pruned, trunk- Mean fruit DM increased from 15.390.13%
girdled and control kiwifruit vines as used to calculate to 19.590.16% over the 7 week period; fruit
hydraulic conductance. B, Hydraulic conductance DM increased at a faster rate in root-pruned
for the soil to shoot pathway (Kshoot; kg MPa 1 s 1 and trunk-girdled vines (Fig. 5). Four and
104). Treatments were applied on 12 February 2009
(dashed line). Data are means n3 vines (n2 for 5 weeks post-treatment, fruit DM content was
root-pruned), 1SEM. significantly increased in trunk-girdled vines
in comparison with control vines. Neither
resulting in consistent and often significant treatment had a visible effect on plant health
reductions. In trunk-girdled vines, ci was sig- or leaf area.
nificantly reduced; however, the response was
delayed in relation to effects on A and gs, with
significant differences emerging 25 days post- Discussion
treatment (Fig. 2C). Net photosynthesis and gs Physiological responses to trunk girdling and
were reduced in root-pruned vines compared root pruning differed markedly in these kiwifruit

Table 2 Post-treatment hydraulic conductance (K; kg MPa 1 s 1104) and kiwifruit scion sapwood area
specific hydraulic conductance (Ks; kg MPa 1 m2 s 1 104). Data are means n 3 vines at six timings (n 2
vines for root-pruned), 91SEM. Values in the same column sharing the same letter are not significantly
different (P B0.05).

Kshoot Kplant Ks-shoot Ks-plant

(kg MPa 1 s 1 104) (kg MPa 1 m2 s 1 104)

Control 6.6290.52 a 3.9790.27 a 0.5790.07 a 0.3090.03 a

Root pruned 4.8990.37 b 3.2690.18 a 0.3990.02 b 0.2590.01 a
Trunk girdled 5.3890.42 ab 3.6390.24 a 0.4790.04 ab 0.3090.02 a
P value 0.0152 0.1655 0.0392 0.3485
Vine responses to girdling and root pruning 37

Figure 4 Fruit fresh weight (g) from fresh weight

calculations following measurements of fruit length
and maximum and minimum diameter at the equa-
torial region of marked kiwifruit for root-pruned,
trunk-girdled and control vines. Treatments were
applied on 12 February 2009 (dashed line). Data are
means n 18 fruit (n12 for root-pruned),
Figure 2 Leaf gas exchange characteristics of root- 1SEM.
pruned, trunk-girdled and control kiwifruit vines. A,
Photosynthetic rate (mmol CO2 m 2 s 1). B, Sto-
matal conductance (mol H2O m 2 s 1). C, Leaf observations in grape (Smart et al. 2006). There
internal CO2 content (ci; mmol CO2 mol 1 air). was evidence of lower A; however, differences
Treatments were applied on 12 February 2009 were rarely significant and were smaller than
(dashed line). Data are means n 612 leaves, expected. The reduction in Kshoot (Fig. 1B) was in
1SEM. * indicates a significant difference in
comparison with the control at that time. line with what might be expected from removing
50% of the root system. In control vines, the soil
vines. The physiological response to root prun- to shoot pathway accounted for 52% of whole
ing demonstrated a primary hydraulic response,
with significant reductions in C shoot, C leaf and
Kshoot. Reductions in C leaf agreed with previous

Figure 5 Fruit dry matter content (% of fruit fresh

weight) of periodically sampled fruit from root-
pruned, trunk-girdled and control kiwifruit vines.
Figure 3 d13C discrimination () at weekly intervals Treatments were applied on 12 February 2009
post-treatment for root-pruned, trunk-girdled and (dashed line). Data are means n 12 fruit (n 8 for
control kiwifruit vines. Data are means n 6 sam- root-pruned), 1SEM. * indicates a significant
ples, 1SEM. * indicates a significant difference in difference in comparison with the control at that
comparison with the control at that time. time.
38 MZ Black et al.

plant hydraulic resistance (Table 2), and the root root-pruned vines (Fig. 2). Consistent, signifi-
system accounts for a large proportion of the cant reductions in gs without consistent reduc-
resistance along this pathway (McAneney & tions in C shoot and C leaf suggest that the
Judd 1983). Assuming the majority of this stomatal response is not hydraulic. This agrees
resistance resides in the root, removing 50% of with previous suggestions that trunk girdling
root area would be expected to double this does not directly affect water transport (Noel
resistance, resulting in an approximate 34% 1970; Goren et al. 2003), instead supporting the
reduction in Kplant. During the first 5 days hypothesis that stomatal closure is coordinated
post-treatment, root-pruned vine Kplant was with a biochemical down-regulation of A. The
reduced by 22%26%; deviation from the esti- delayed reduction in leaf ci and increase in
mated 34% could be attributed to resistance d13C (Fig. 3) suggests that during the 3 weeks
along the trunk and shoot. Effects on Kshoot post-treatment, A was limited by combined
appeared transitory, with Kshoot similar in root- biochemical down-regulation and stomatal
pruned and control vines by 4 weeks post- limitation, after this time stomatal limitation
treatment. However, this appears to be due to appears to have been dominant. Differences in
reduced K in control vines over time, with root- leaf d13C content between control and girdled
pruned vine Kshoot remaining lower than pre- vines became less evident 5 weeks post-treat-
treatment. ment. Wounds of trunk girdles applied to
Hydraulic conductance and Eplant were low orchard-grown vines in spring and early sum-
in these root bag-grown vines in comparison mer generally callus within 4 weeks, similar to
with observations for orchard-grown kiwifruit the healing time observed in grape (Williams
vines (Judd et al. 1986; Green et al. 1989; et al. 2000). Commencement of photosynthetic
Clearwater et al. 2004). Treatments appeared to recovery at 5 weeks may indicate reconnection
have little effect on whole vine Eplant, even of phloem vasculature at this time.
when gs was reduced. This may be a result of Reduced gs and A following trunk girdling
environmental conditions within the plastic have been observed previously (Williams et al.
greenhouse, with restricted air movement redu- 2000) and the role of feedback inhibition of A
cing stomatal control of transpiration (Jarvis & following reduced photosynthate export from
McNaughton 1986). leaves has been proposed (Iglesias et al. 2002).
Root pruning was expected to have a Trunk girdling temporarily removes the root
significant effect on photosynthesis, through system from the source:sink ratio, increasing
stomatal limitation of leaf gas exchange. The availability of photosynthates to parts above
lack of a consistent effect on gs in kiwifruit the girdle. A proportion of these newly avail-
conflicts with previous findings in other species. able photosynthates may be utilized for growth;
In grape, Smart et al. (2006) observed signifi- however, girdling has been shown to result in
cant reductions in gs that persisted for at least accumulation of carbohydrates within leaves,
2 months. In the present experiment, reductions reducing gs and A (Iglesias et al. 2002; Domec
in C leaf without significant reductions in gs & Pruyn 2008). In contrast, Williams et al.
suggest that kiwifruit vines are anisohydric (2000) suggested that the accumulation of
(Tardieu & Simonneau 1998) and that reduc- carbohydrates in girdled grape vine leaves was
tions in C leaf observed were insufficient to insufficient to result in feedback inhibition of
prompt a stomatal response. The lack of a A and suggested the involvement of abscisic
significant effect on ci or d13C following root acid (ABA) effects on stomatal aperture. This
pruning supports the observation of only minor hypothesis is supported by previous observa-
reductions in gs. tions of increased ABA accumulation in leaves,
The photosynthetic response to trunk coinciding with stomatal closure following
girdling was more pronounced than that in girdling (Düring 1978; Setter et al. 1980).
Vine responses to girdling and root pruning 39

Fruit fresh weight and growth rate were not explaining the variable results of root pruning
affected by either treatment. Root pruning in on fruit quality parameters in previous studies.
other fruit crops has resulted in negligible
or negative effects on fruit growth, with
reductions in fruit size and overall yield †
Funding was provided through a ZESPRI Group
(Khan et al. 1998; Ferree et al. 2000). Previous
Ltd-funded PhD scholarship as part of the ‘Taste
observations of kiwifruit fresh weights follow-
Improvements via Kiwifruit Root Manipulation’
ing root pruning have been variable (Patterson project TZ0847 and The New Zealand Foundation
et al. 2009). Trunk girdling was expected to for Research, Science and Technology project
increase kiwifruit fresh weights following C06X0706. Peter Minchin, Bill Snelgar and Steve
previous observations (Currie et al. 2005). Green are acknowledged for their advice and assis-
Increases in fruit DM have previously been tance with equipment.
obtained from both treatments (Currie et al.
2005; Patterson et al. 2009) and trunk girdling
results from this experiment concur. In-
Clearwater MJ, Lowe RG, Hofstee BJ, Barclay C,
consistent effects on fruit DM following root
Mandemaker AJ, Blattmann P 2004. Hydraulic
pruning in the present experiment suggest that conductance and rootstock effects in grafted
the technique has the potential to increase DM; vines of kiwifruit. Journal of Experimental
however, timing and severity may be important Botany 55: 13711382.
considerations. Currie MB, Barnett AM, Boyd LM, Max S 2005.
In summary, hydraulic and photosynthetic Trunk girdling: risks and opportunities. New
Zealand Kiwifruit Journal 167: 1217.
responses of kiwifruit vines were investigated in Currie MB, Jackman R, Max S, Blattmann P,
response to trunk girdling and root pruning. Seymour S 2008. Summer girdling*current
These two practices can result in increased fruit options and new ideas. New Zealand Kiwifruit
size and quality, so it was of interest to see how Journal 185: 1317.
they compared in their physiological effects. De Schepper V, Steppe K, Van Labeke M, Lemeur R
2010. Detailed analysis of double girdling effects
The observed responses differed markedly. The
on stem diameter variations and sap flow in
response to root pruning was hydraulic, with young oak trees. Environmental and Experi-
significant reductions in C and K, supporting mental Botany 68: 149156.
our hypothesis. Photosynthetic responses Detling JK, Winn DT, Procter-Gregg C, Painter EL
following root pruning were less severe than 1980. Effects of simulated grazing by below-
expected, attributed to kiwifruit vines demon- ground herbivores on growth, CO2 exchange,
and carbon allocation patterns of Bouteloua
strating an anisohydric stomatal response. gracilis. Journal of Applied Ecology 17: 771
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