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Methods to Minimize the Effect of Ethylene Sprout Inhibitor on Potato Fry

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Article  in  European Potato Journal · December 2006

DOI: 10.1007/s11540-007-9025-6

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Potato Research (2007) 49:303–326
DOI 10.1007/s11540-007-9025-6

Methods to Minimize the Effect of Ethylene Sprout

Inhibitor on Potato Fry Colour

Barbara J. Daniels-Lake & Robert K. Prange &

Wilhelmina Kalt & John R. Walsh

Received: 29 August 2006 / Accepted: 30 July 2007 /

Published online: 28 September 2007
# EAPR 2007

Abstract Ethylene is an effective potato (Solanum tuberosum L.) sprout inhibitor,

but it often darkens fry colour. Trials were conducted over nine consecutive storage
seasons to identify ethylene application methods which would mitigate darkening
while retaining adequate sprout inhibition, using cv. Russet Burbank plus cvs
Shepody, Asterix and Santana in some years. Tubers were stored for up to 35 weeks in
closed chambers with ethylene gas delivered via the ventilation airstream. Exposure to
continuous 4 μl l−1 ethylene after suberization and cooling were completed was
designated the ethylene check. Alternative ethylene treatments included commencing
exposure either before or after suberization was completed; gradually introducing the
ethylene either by a concentration gradient of eight steps over 4 or 8 weeks or by
increasing the duration of exposure from 6 to 24 h per day in four weekly steps;
repeatedly interrupting exposure for several hours per day or for durations of 1 or
more days; and warm storage. Selected ethylene treatment combinations were applied
in each year, plus untreated controls, chlorpropham-treated (CIPC) checks and
ethylene checks. Sprout growth, fry colour, loss of mass and disease incidence were
evaluated at regular intervals. In all cultivars and all years, the ethylene check
darkened fry colour more than the other treatments. Commencing before suberization
ended, gradually introducing the ethylene by either concentration or time gradient and
interrupting the exposure all reduced the negative effect of ethylene sprout inhibitor on
fry colour. Continuous ethylene treatments inhibited sprout growth as effectively as
CIPC, except at 13 °C storage. Interruptions of 18 h and 2 or more days reduced
sprout inhibition. Regardless of cultivar variations, an early start using either a
concentration or time-increment gradient had the least effect on fry colour while
maintaining good sprout inhibition.

B. J. Daniels-Lake (*) : R. K. Prange : W. Kalt

Atlantic Food and Horticulture Research Centre, Agriculture and Agri-Food Canada,
Kentville, NS B4N 1J5, Canada
e-mail: danielslakeb@agr.gc.ca

J. R. Walsh
McCain Foods Limited, Florenceville, NB E7L 3G6, Canada
304 Potato Research (2007) 49:303–326

Keywords Chlorpropham . Postharvest . Solanum tuberosum . Sprout inhibition .

Storage

Abbreviations
ARu Agtron reflectance units, % reflectance
Ast cv. Asterix
conc Concentration gradient
post-sub After suberization
PH Postharvest
pre-sub Before suberization
RB cv. Russet Burbank
San cv. Santana
Shp cv. Shepody

Introduction

In recent years, several alternatives to the widely used potato sprout inhibitor
chlorpropham [CIPC; isopropyl (n-3-chlorophenyl) carbamate] have been identified
and developed. Among these, ethylene gas has been registered for several years in
Canada and the United Kingdom, and is currently used on approximately 50,000 t of
table stock annually in England (J. Barnes, BioFresh Inc., personal communication).
Ethylene effectively controls sprouting for up to 30 weeks, is relatively inexpensive
to apply, and leaves no residue (Prange et al. 1998). However, ethylene also increases
the respiration rate of the tubers and promotes conversion of starch to sugars, potentially
increasing the sugar concentrations in treated tubers (Reid and Pratt 1972; Day et al.
1978; Prange et al. 1998). High reducing sugar concentrations (predominantly glucose
and fructose) can lead to dark fry colour when the tubers are processed into potato
chips or French fries. Since fry colour is of paramount importance in the potato
processing industry, processors are reluctant to adopt ethylene sprout inhibition and
thereby risk reducing the quality of their processed products.
After a suberization period and gradual cooling to long-term storage temperature
(9 °C), Prange et al. (1998) applied 4 μl l−1 ethylene continuously for sprout
inhibition, from early December to the end of the storage season, i.e. beginning
about 8–10 weeks after harvest. Fry colour using this ethylene application method is
darker than with CIPC sprout inhibitor in some storage years. The fry colour becomes
progressively lighter during continued storage with ethylene, although it does not fully
recover to the colour of check tubers (Prange et al. 1998). Reconditioning, i.e. warm
storage prior to processing to reduce sugar concentrations, also improves the fry
colour of potatoes affected by ethylene sprout inhibitor, but the colour is seldom as
light as CIPC-treated tubers (Prange et al. 2005a).
Daniels-Lake et al. (2005) found that ethylene concentrations below 4 μl l−1 have
less effect on both fry colour and sprout growth of cv. Russet Burbank. In contrast,
40 and 400 μl l−1 ethylene does not cause increased darkening compared to 4 μl l−1,
but provides marginally better sprout inhibition. This suggests that different
metabolic pathways are involved in the sweetening and sprout-inhibitory effects of
Potato Research (2007) 49:303–326 305

ethylene, which implies that the two can be manipulated separately. Short or
intermittent exposure to ethylene has been shown to promote sprouting (Pratt and
Goeschl 1969; Rylski et al. 1974; Timm et al. 1986). However, the effects of further
variations in ethylene application methodology have not been well described.
Studies were undertaken to determine whether manipulating storage temperature,
start date, continuity of exposure, initial ethylene concentration, and initial exposure
period, or combinations of these methods, would reduce the negative effect of
ethylene sprout inhibitor on fry colour while maintaining good sprout inhibition.

Materials and Methods

Trials were conducted at the Agriculture and Agri-Food Canada (AAFC) postharvest
research facilities at the Atlantic Food and Horticulture Research Centre (AFHRC)
in Kentville, Nova Scotia, Canada, over nine consecutive fall to spring storage
seasons, i.e. 1991–1992 through 1999–2000 inclusive.
Each fall, potato tubers from four commercial growers were obtained from
McCain Foods, Florenceville, New Brunswick, Canada and/or Cavendish Farms,
Summerside, Prince Edward Island, Canada. In 1997–1998 only, one of the four
grower-lots of potatoes of each cultivar tested was obtained from a commercial
producer in the province of Manitoba, Canada. Most of the trials used cv. Russet
Burbank (RB) potatoes, with cv. Shepody (Shp) potatoes used in some. Both are
grown extensively in North America for French fry processing. Cvs Asterix (Ast)
and Santana (San) potatoes were also utilized in one trial-year.
Within each October to June storage season, four lots of RB or Shp tubers
constituted the four replicates used in the trial; replication of Ast and San was from
within a single lot of tubers for each cultivar. Potatoes were sourced from different
growers each year to ensure that the tubers tested were representative of the
commercially available material. Tubers for the 1991–1992 and 1992–1993 seasons
were suberized after harvest in a commercial potato storage facility (3–4 weeks at
approximately 13 °C in darkness), before shipping to AFHRC. The tubers were then
stored in darkness, cooled at a rate of 1°C per week to 9 °C, and held at 9 °C for 2–
4 weeks until the start of the trials in early December of each year. In the 1994–1995
and subsequent storage seasons, potatoes were shipped to AFHRC in early October
shortly after harvest and the suberization and cooling phases of storage were
incorporated into the storage trials. These tubers were initially held in darkness at
13 °C for 3–4 weeks and then cooled as above.
Samples of healthy, reasonably uniform tubers (9–11 tubers, 150–300 g each,
total sample mass c. 2 kg) were selected from within each of the four grower-lots of
potatoes and placed in small onion-mesh bags. Samples were assigned to each of the
treatments and evaluation dates of the trial according to the experimental design. The
samples were labeled, weighed and stored in sealed storage chambers. From 1991–
1992 through 1994–1995 the chambers were 64-l PVC barrels (IPL, Moncton, New
Brunswick, Canada), mounted horizontally in a wooden rack. However, these barrels
deteriorated considerably over time and new, sturdier 0.34 m3 stainless steel
chambers with acrylic lids were constructed locally and used in the 1995–1996
through 1999-2000 seasons. Each barrel initially contained five samples from a
 306

Table 1 Ethylene treatment methods and cultivars tested in each storage season

Treatment Year
91–92 92–93 93–94 94–95 95–96 96–97a 97–98 98–99 99–00b

a. Long-term storage at 9 °C after suberization and gradual cooling

Control: untreated; suberized, cooled and stored without sprout inhibitors
RBc RB, Shpd RB RB RB RB RB, Shp RB, Shp, Ast, San RB
CIPC check: chlorprophame dip after suberization and cooling were completed
RB RB, Shp RB RB RB RB RB, Shp RB, Shp, Ast, San RB
Ethylene: continuous 4 μl l−1, except as modified by the treatments below
f
Start time: pre-sub
Abruptg –h – – RB RB RB RB, Shp RB, Shp, Ast, San RB
Gradual: Conci – – – – – RB RB, Shp RB, Shp, Ast, San RB
j
Conc–fast – – – – – RB – – –
Timedk – – – – – – – – RB
Interrupted (hours):l 18 h – – – – RB – – – –
12 h – – – – RB – – – –
6h – – – – RB – – – –
Start time: Post-subm
Abrupt – – – – – RB – – –
Gradual: Conc–fast – – – – – RB – – –
n
Start time: Cooled
Abrupt (eth checko) RB RB, Shp RB RB RB RB RB, Shp RB, Shp, Ast, San RB
Gradual: conc – – – – – – RB, Shp – –
Interrupted (hours): 6 h – – – – RB – – – –
12 h – – – – RB – – – –
18 h RB RB
Potato Research (2007) 49:303–326

– – – – – – –
Table 1 (continued)

Treatment Year
91–92 92–93 93–94 94–95 95–96 96–97a 97–98 98–99 99–00b

Interrupted (days):p 1 day – RB – – – – – – –

2 days – RB – – – – – – –
3 days – RB – – – – – – –
6 days RB – – – – – – – –
b. Long-term storage at 13 °C, after suberization, cooling to 9 °C and re-warming in mid-January
Control – – – – – – – Shp RB
CIPC check – – – – – – – – RB
Ethylene (4 μl l-1)
Start time: Mid-January
Abrupt – – – – – – – Shp RB
Gradual: Conc – – – – – – – Shp –
Timed – – – – – – – Shp RB
Potato Research (2007) 49:303–326

a
October to mid-January only; samples were assessed at intervals of 2 weeks
b
Evaluations in 1999–2000 were not at regular 5-week intervals, but were less frequent during the winter and more frequent during the spring
c
Cultivars tested: RB cv. Russet Burbank, Shp cv. Shepody, Ast cv. Asterix, San cv. Santana
d
Shepody was stored until late March only; samples were assessed at intervals of 3 weeks
e
Dipped in a 1% emulsion of isopropyl (n,3-chlorophenyl) carbamate in early December
f
Pre-suberization; ethylene exposure commenced as soon as possible after harvest (1–5 weeks postharvest, ca. mid-October), i.e. before suberization was finished
g
Abrupt start of ethylene exposure, i.e. 0–4 μl l−1 within a few hours
h
Dash indicates that the treatment was not applied in that year
i
Gradual ethylene exposure by concentration gradient; ethylene exposure commenced at a very low concentration with weekly increments to 4 μl l−1
j
Concentration gradient: fast; rapidly increasing the ethylene concentration by changing at half-weekly intervals
k
Gradual ethylene exposure by timing changes; exposure to continuous 4 μl l−1 ethylene for 6 h per day in the first week of storage, 12 h per day in week two, 18 h per day in
week three and 24 h per day thereafter
l
Ethylene exposure was discontinued for a period of x h each day
m
Post-suberization; ethylene exposure commenced after the suberization period ended but before cooling to the long-term storage temperature (5–9 weeks postharvest, c. mid-
November)
n
Post-cooling; ethylene exposure commenced after completion of suberization and cooling to the long-term storage temperature (9–12 weeks postharvest, ca. early December)
o
Ethylene check treatment; abrupt start of continuous exposure to 4 μl l−1 , after cooling to 9 °C for long-term storage
p
Ethylene exposure was discontinued for the indicated number of days after 1 day of exposure; this cycle was repeated until the end of the storage term
307
308 Potato Research (2007) 49:303–326

single grower, and each treatment was applied to four barrels. Each stainless steel
chamber contained four PVC baskets (NPL-655; Norseman Plastics, Rexdale,
Ontario, Canada) which each held five to eight samples from a single grower, and
each treatment was applied to one stainless steel chamber. All chambers were stored
inside a refrigerated coldroom for temperature control.
Humidified ventilation air was supplied to the chambers via 3 mm i.d. nylon
tubing (Bowman Products, Moncton, New Brunswick, Canada) which entered
through one end of each chamber and was exhausted at the opposite end. Exhaust air
was drawn outside the building by an exhaust fan. Ventilation air was supplied to
each PVC barrel at c. 0.5 l min−1 for 6 h per day (ventilation period), followed by
18 h per day without ventilation (static period) during 1991–1992 and 1992–1993.
Due to concern about possible CO2 accumulation and O2 depletion during the 18-
h static periods, ventilation was increased to two 6-h ventilation periods alternated
with 6-h static periods each day during 1993–1994. The stainless steel chambers
were ventilated at c. 2 l min−1 for two 6-h ventilation periods, separated with 6-h
static periods, each day. These ventilation rates were lower than the rates used in
commercial potato storage facilities, where ventilation is used to manage pile
temperature. In this study, ventilation served to deliver the ethylene treatments and to
maintain O2 and CO2 at near-ambient concentrations, but not to control temperature.
Therefore these ventilation rates should not be compared directly with rates used in
commercial settings.
Sufficient ethylene from a compressed-gas cylinder (Canadian Liquid Air,
Kentville, Nova Scotia, Canada) was added to the ventilation air, at a gas
distribution board constructed on-site, to provide the desired ethylene concentration
in the chambers. The ventilation and ethylene delivery schedules were controlled by
multi-channel timers (ChronTrol, San Diego, California). Ethylene concentration in
the chambers was quantified by gas chromatography, using the method of Prange et
al. (1998). Any necessary adjustments to maintain the desired concentrations ± 10%
were made by manually adjusting needle valves (Nupro, Swagelok Company, Solon,
Ohio) on the gas distribution board. Rapid increases in ethylene concentration within
a chamber were achieved by sharply increasing the flow of ethylene gas added to the
ventilation airstream of the chamber for several seconds at the beginning or end of a
ventilation cycle. The added gas dispersed throughout the chamber within 10–
20 min. Rapid decreases in ethylene concentration were achieved by increasing the
ventilation flow-rate to all chambers to c. 15 l min−1, for up to 30 min, at the
beginning or end of a regular ventilation cycle, flushing the ethylene-air mixture in
the chamber to exhaust. Additional details regarding the treatment chambers and
associated apparatus can be found in Daniels-Lake (2001).
The method of Prange et al. (1998), whereby tubers stored at 9 °C were exposed
to 4 μl l−1 ethylene gas continuously from early December (after suberization and
cooling) until June, was designated the ethylene check (eth check). The alternative
ethylene application methodologies explored in this work and the cultivars tested are
described in Table 1. Figure 1 depicts the temperature regime applied in all years,
and also the changes in ethylene concentration with time in the various treatment
regimes (except the interrupted methods).
During each successive storage season, one or more of the ethylene methodol-
ogies was included and the responses compared with the eth checks, untreated
Potato Research (2007) 49:303–326 309

Fig. 1 Temperature and ethylene concentration profiles

controls and CIPC checks (except no CIPC check in 1996–1997). Untreated control
samples received no sprout inhibitor treatment; storage and ventilation were as
described above. CIPC check samples were dipped in 1% a.i. (v/v) water emulsion of
isopropyl (n-3-chlorophenyl) carbamate (Sprout-Nip EC, Stanchem, Etobicoke,
Ontario, Canada) in early December; storage and ventilation were as described above.
The series of trials progressed through several successive years, seeking an
effective method(s) to apply ethylene as a sprout inhibitor without seriously affecting
fry colour. Finite resources and the practical concerns of the potato industry sponsors
310 Potato Research (2007) 49:303–326

resulted in obviously ineffective methods being promptly discontinued, while

promising methods were repeated and built upon. In this way the selection of
treatments evolved rapidly. Although certain specific treatment methodologies were
tested in only a single year, treatments of a similar type were evaluated in multiple
years and in some cases using two or more cultivars (Table 1). The untreated control,
CIPC check and eth check were employed consistently every year. The eth check in
particular provided a baseline against which responses to the other treatments could
be compared, within the comprehensive context of the research. Clear trends
emerged, which are discussed in the text.
Several tuber responses were evaluated, including number and mass of sprouts
(>5 mm), length of the longest sprout within a sample, fry colour, weight loss, and
disease incidence. Fry colour was assessed as described by Daniels-Lake et al.
(2005), by deep-frying discs cut from a central longitudinal slice from each tuber.
One disc from each of the 10 tubers in a sample was evaluated. The colour of the
cooled discs was measured with an Agtron reflectance colourimeter (M-35-D,
Agtron, Sparks, Nevada, USA). The percent reflectance scores were expressed as
Agtron % reflectance units (ARu), whereby a light-coloured fried disc received a
high numerical score and a dark disc received a low score. Scores greater than 100
ARu were possible, if the colour of the fried disc was lighter than the reference disc
(Agtron # 56 reflectance calibration disc, very pale grey in colour; Agtron, Sparks,
Nevada, USA) used for calibration of the instrument. Agtron values of 89–100 ARu
are approximately equivalent to USDA color grade #1, 70–88 ARu to USDA grade
#2 and 54–69 ARu to USDA grade #3.
Dry matter was measured as described in Daniels-Lake et al. (2005). The affected
portion of any diseased tubers was visually estimated as a fraction of the total
surface area of the tuber. The occurrence of tuber disorders such as brown centre-
hollow heart, tuber greening, vascular discolouration or any other disorders, was
recorded when observed.
Evaluations were conducted at the start of each storage trial (fry colour and dry
matter only), after the suberization period (1994–1995 and later trials), after cooling
(1994–1995 and later trials), and at 5-week (RB, Ast, San) or 3-week (Shp) intervals
thereafter to the end of each storage season, except as noted otherwise. In the 1996–
1997 trial only, evaluations were conducted at the start of the trial, i.e. 1 week
postharvest (PH), 1 week later (2 weeks PH), and then bi-weekly until 14 weeks PH,
except that there was no evaluation at 12 weeks PH. Because RB has a long
dormancy period and does not usually sprout until February or March, there were no
sprout data from the 1996–1997 trial which ended in mid-January 1997. For the
same reason, a CIPC check was not included in the 1996–1997 trial.
Each storage season (year) comprised a trial, using a randomized split-split plot
design. Treatment was the main plot, grower was the sub-plot and evaluation date was the
sub-sub-plot. The data from each trial were evaluated by analysis of variance (ANOVA),
using Genstat statistical software (Payne 2005). For all characteristics evaluated, data
from several trial years were combined in multi-year analyses whenever possible.
Sprout data were mathematically transformed to log10 values prior to statistical
analysis, to normalize the data. To counteract the biasing effect of numerous zeros in
the sprout data, the CIPC treatment and the early evaluation dates before sprouting
commenced (i.e. 0-values) were excluded from the statistical analyses. When
Potato Research (2007) 49:303–326 311

significant differences were identified (P≤0.05), means were compared using the
least significant difference. Statistical comparisons among the transformed means
were made before the values were back-transformed for presentation and discussion.
Unless otherwise noted, only results significant at P≤0.05 are discussed.

Results

Throughout these trials, fry colour of ethylene-treated tubers darkened soon after the
exposure commenced and usually recovered, as observed by Prange et al. (1998) and
Daniels-Lake (2001). However the degree of darkening and recovery varied among
the treatments, as described below. Each year, sprout growth increased with time
during the storage term; tubers progressively broke dormancy and the sprouts
elongated. A delay in the first appearance of measurable sprouts (>2 mm) and weak
attachment of sprouts, as described by Prange et al. (1998), were also observed in
these trials (data not presented). Since sprout growth naturally increases with time,
only sprout data from the end of the trial (i.e. the maximum sprout development) are
discussed.
In tubers stored at 9 °C, there were no differences in dry matter or disease
incidence attributable to treatments (data not presented). The loss of tuber mass
increased slightly at each successive evaluation date, but there were no differences
associated with treatments in any trials stored at 9 °C, except for the untreated
controls. In most years, loss of mass in the control tubers was greater than in any
other treatments (data not presented). This was attributable to the heavy sprout
development observed in the later stages of the trials, and to the accompanying res-
piration and transpiration losses from the sprouting tubers. In tubers stored at 13 °C,
disease development and weight loss were higher than in tubers stored at 9 °C (c.f.,
section “Elevated Storage Temperature”).

Interrupted Exposure (Days)

Exposure of RB tubers for 1 day followed by 6 days without ethylene (1991–1992

season) did not darken tuber fry colour in comparison with the CIPC check (e.g. 68
and 70 ARu, respectively, at 24 weeks PH), but sprout inhibition was inadequate
(data not presented). In the following year (1992–1993) the duration of interruptions
between ethylene exposure-days was reduced to 3, 2, or 1 day. The fry colour of
tubers in the ethylene treatment interrupted for 3-day intervals was lighter than in the
CIPC check and eth check tubers at 13 and 23 weeks, and was lighter than control
tubers at 33 weeks PH (Table 2a). At all evaluations, fry colour in the ethylene
treatments interrupted for 1- or 2-day intervals was not significantly different from
the CIPC check, eth check and control tubers. Surprisingly, fry colour of tubers in
the eth check was also not different from the control or CIPC check tubers at any
evaluation. However, it should be noted that the initial fry colour of all four lots of
potatoes used in that year was rather dark (mean 50±2.63 ARu), and remained so
throughout the trial (Table 2a).
In contrast to the fry colour observations, sprouting was clearly affected by the
ethylene treatments. Sprout elongation was inhibited more strongly in the eth check
Table 2 Effects of interrupted ethylene exposure on potato fry colour and sprouting, cv. Russet Burbank
312

Treatment Fry coloura (Agtron % reflectance) Maximum sprout Sprout massb, 33 weeks (g kg−1)
c
lengthb, 33 weeks (mm)
13 weeks 18 weeks 23 weeks 28 weeks 33 weeks

a. Interruptions of 1, 2, or 3 days’ duration, 1992–1993

Untreated control 48.9 abd 52.3 52.6 ab 54.9 51.3 b 35.7 (1.565 a) 3.3 (0.601 a)
CIPC check 47.0 b 52.2 50.5 b 56.1 54.6 ab 0.3 (0.119 c) 0.0 (0.0 b)
Ethylenee
Eth check 49.1 b 52.0 50.7 b 52.3 55.4 ab 0.7 (0.226 c) 0.0 (0.001 b)
Interrupted exposure
1 day 46.1 b 49.9 52.2 ab 53.2 54.6 ab 0.8 (0.260 c) 0.0 (0.014 b)
2 days 49.4 ab 49.7 53.6 ab 52.7 53.5 ab 4.0 (0.703 bc) 0.7 (0.234 b)
3 days 52.2 a 49.8 nsf 55.0 a 54.1 ns 56.2 a 10.1 (1.044 ab) 0.3 (0.103 b)
SE = 1.387 0.2110g 0.003
P= 0.046 0.001 0.003

b. Interruptions of several hours each day, 1995–1996

Untreated control 82.7 a 83.9 a 83.2 a 84.7 a 81.1 a 665 (2.824 a) 60.1 (1.7861 a)
CIPC check 80.7 a 85.9 a 84.0 a 87.5 a 85.2 a 0.0 (0.000 d) 0.0 (0.000 d)
Ethylene
Eth check 55.2 c 64.2 c 67.0 c 74.6 b 73.4 b 27 (1.452 c) 0.2 (0.0825 d)
Interrupted exposure
6h 68.1 b 72.8 b 76.3 b 78.5 b 82.1 a 60 (1.785 c) 11.9 (1.1090 c)
12 h 68.9 b 76.3 b 80.1 ab 85.6 a 82.7 a 229 (2.362 b) 27.8 (1.4588 b)
18 h 72.2 b 82.9 a 81.9 ab 85.4 a 85.2 a 510 (2.708 ab) 50.3 (1.7103 a)
SE = 2.030 0.393 0.03138
P= <0.001 <0.001 <0.001

a
Initial fry colour (8 weeks postharvest) was 50.0±2.63 and 82.3±3.18 Agtron % reflectance units in 1992–1993 and 1995–1996, respectively
b
Back-transformed values; log10 means from the statistical analyses are in parentheses
c
Weeks after tubers were harvested, treatment × evaluation date interaction
d
For each year, values within a column followed by the same letter are not significantly different at P≤0.05. To maintain clarity of the fry colour data, differences across evaluation
dates have not been indicated. For these comparisons, three times the SE can be used to closely approximate the least significant difference between means
e
Ethylene treatments are as described in Table 1
f
Not significantly different at P≤0.05
g
Potato Research (2007) 49:303–326

SE and P values for sprouting characteristics pertain to log10 values used in the statistical analysis
Potato Research (2007) 49:303–326 313

(in which ethylene exposure occurred daily) than when exposure was interrupted for
3-day intervals (Table 2a). At 33 weeks PH, the maximum length of sprouts on
tubers in the eth check and the ethylene treatments interrupted for 1- or 2-day
durations was not different from tubers in the CIPC check. Sprouts in all ethylene
treatments interrupted for 1 or 2 days were shorter than sprouts on control tubers.
However, when the ethylene was interrupted for a duration of 3 days, tuber
maximum sprout length was not significantly different from that of untreated control
tubers. The mass of sprouts at 33 weeks PH in all ethylene treatments was not
significantly different from the CIPC check, and was much lower than on the control
tubers (Table 2a).

Interrupted Exposure (Hours)

Fry colour was darkest in the eth check at all evaluation dates (Table 2b).
Interruption of the ethylene exposure for 6, 12 or 18 h per day reduced the darkening
of tuber fry colour compared with the eth check (in which ethylene exposure was
continuous for 24 h per day). The exception to this was at the 28-week evaluation
when the fry colour of tubers in the 6-h interruption treatment was not significantly
different from the eth check (Table 2b).
At the first evaluation (13 weeks PH) fry colour in all ethylene treatments
was darker than in the CIPC treatment, but it recovered (lightened)
progressively at subsequent evaluations (Table 2b). Fry colour in the eth check
remained darker than in the CIPC check throughout the trial (Table 2b). In the
ethylene treatments interrupted for 18, 12 and 6 h per day, fry colour improved
sufficiently to match the colour in the CIPC check treatment, beginning at the 18,
23 and 33 week evaluations, respectively. Fry colour of the untreated control
tubers and the CIPC-treated check tubers was not different at any evaluation date
(Table 2b).
Maximum sprout length and sprout mass of tubers in the 18-h interruption
treatment were not different from the control tubers at 33 weeks PH, but were
greater than CIPC-treated tubers (e.g. lengths of 510, 665 and 0 mm,
respectively; Table 2b). Although the maximum sprout length of tubers exposed
to the eth check was longer than of tubers treated with CIPC, sprouts were much
shorter than on control tubers (27, 0 and 665 mm, respectively, at 33 weeks PH).
Sprout mass in the eth check was not significantly different from CIPC-treated
tubers (Table 2b). Mass and length in the 12- and 6-h interruption treatments were
intermediate between the 18-h interruption and eth check (continuous ethylene)
treatments.

Elevated Storage Temperature

In Shp in 1998–1999, the fry colour of tubers stored with ethylene at 13 °C was
lighter than tubers in the eth check stored at 9 °C (Table 3a). Tubers stored with ethylene
at 13 °C had fry colour as light as CIPC check tubers stored at 9 °C (Table 3a).
Maximum length, sprout mass and weight loss in the ethylene treatments stored at
13 °C were all greater than CIPC-treated tubers stored at 9 °C, but were less than in
the controls stored at either temperature. However, among the ethylene treatments,
314 Potato Research (2007) 49:303–326

Table 3 Effects of increasing storage temperature to 13 °C vs. continuous storage at 9 °C on potato fry
colour, sprouting and weight loss

Treatment Fry coloura Weight lossb Maximum Sprout massb

(Agtron % reflectance) (%) sprout (g kg−1)
length (mm)

a. cv. Shepody, 1998–1999 All evaluationsc 26 weeksd 26 weeksd 26 weeksd

Untreated control,
9 °C 97.3 abe 5.2 a 148.6f (2.175 b) 29.0f (1.4773 b)
13 °C 101.5 a 5.9 a 379.2 (2.580 a) 79.5 (1.9058 a)
CIPC check, 9 °C 93.3 bc 2.5 c 0.0 (0.000 d) 0.0 (0.0000 e)
Ethyleneg
Eth check (9 °C) 67.4 d 3.5 b 17.0 (1.256 c) 2.3 (0.4953 d)
mid-Jan abrupt, 88.4 c 4.0 b 23.3 (1.385 c) 11.2 (1.0879 c)
13 °C
mid-Jan conc, 93.8 b 3.7 b 26.2 (1.434 c) 12.8 (1.1318 c)
13 °C
mid-Jan timed, 95.7 b 3.9 b 23.3 (1.385 c) 12.2 (1.1220 c)
13 °C
SE = 1.633 0.2816 0.1042h 0.05438h
P= < 0.001 < 0.001 < 0.001 < 0.001

b. cv. Russet Burbank, All evaluationsi All All evaluationsi 35 weeksd

1999–2000 evaluationsi

Untreated control,
9 °C 80.8 8.3 94.2 (1.974) 52.3 (1.727 a)
13 °C 79.9 10.2 170.4 (2.234) 68.3 (1.841 a)
CIPC check, 79.9 7.8 0.0 (0.000) 0.0 (0.000 c)
9 °C
13 °C 82.6 9.5 0.0 (0.000) 0.0 (0.000 c)
g
Ethylene
Eth check (9 °C) 68.7 8.6 3.9 (0.687) 0.1 (0.039 c)
mid-Jan abrupt, 74.8 11.9 14.1 (1.180) 7.4 (0.923 b)
13 °C
mid-Jan timed, 76.0 7.8 7.8 (0.943) 0.6 (0.207 c)
9 °C
13 °C 75.8 9.1 25.1 (1.417) 9.7 (1.029 b)
nsj ns ns
SE = 3.074 0.612 0.1273 0.1326
P= 0.662 0.399 0.606 <0.001

a
Initial fry colours at the start of the trials were 100.1±3.41 and 80.5±8.14 Agtron % reflectance units in
Shepody (17 weeks postharvest) and Russet Burbank (14 weeks postharvest), respectively
b
Proportion of the initial tuber mass at the start of this trial
c
Mean of all evaluations during the trial; significant differences in treatment main effect (P≤0.05)
d
Weeks after tubers were harvested, treatment × evaluation date interaction
e
For each cultivar, values within a column followed by the same letter are not significantly different at
P≤0.05
f
Back-transformed values; log10 means from the statistical analysis are in parentheses
g
Ethylene treatments as described in Table 1
h
SE and P values for sprouting characteristics pertain to log10 values used in the statistical analysis
i
Mean of all evaluations, treatment × temperature interaction
j
Not significantly different at P≤0.005
Potato Research (2007) 49:303–326 315

sprout mass was greater at 13 °C than at 9 °C (Table 3a). Somewhat more disease was
observed among the tubers stored at 13 °C than at 9 °C regardless of treatment, which
is not surprising in view of generally accepted effects of temperature on disease
development. However, insufficient data were available for statistical analysis and
therefore additional research would be needed to confirm or disprove this observation.
In RB tubers tested during the following year, the effects on fry colour, weight loss
and maximum sprout length were similar to the observations in Shp (Table 3b). The
darkest fry colour was observed in the eth check treatment (which was stored at 9 °C),
while the other treatments had lighter fry colour, regardless of storage temperature.
However, the treatment × temperature interaction was not significant (P=0.662; Table
3b). The main effect of treatment on fry colour was significant (P<0.001), regardless
of temperature; i.e. the CIPC checks had the lightest colour, followed by the controls.
The darkest colour was observed in the abrupt ethylene treatments.
The main effects of both treatment and temperature on weight loss in RB during
1999-2000 were significant (P=0.033 and <0.001, respectively). Regardless of
temperature, weight loss was greatest in the abrupt ethylene treatments and least in
the CIPC check, while the controls were intermediate between these (10.2, 8.6 and
9.2% of initial mass, in the abrupt ethylene treatments, CIPC check and controls,
respectively). Mean weight loss in response to temperature was greater in tubers
stored at 13 °C than at 9 °C (10.2 and 8.1% of initial mass, respectively).
Sprout mass of tubers in the eth check (stored at 9 °C) and mid-Jan timed treatment
(stored at 13 °C) was equivalent to the sprout mass of the CIPC check tubers stored at
either temperature (Table 3b). Sprout mass in the ethylene treatment stored at 13 °C
was intermediate between that of ethylene-treated tubers stored at 9 °C and CIPC-
treated tubers stored at either temperature. The observations of maximum sprout length
were similar to the sprout mass findings, but the treatment × temperature interaction was
not significant. However, the main effects of both treatment and temperature on
maximum sprout length were significant (P<0.001 for each). Regardless of
temperature, maximum length was greatest in the controls and least in the CIPC
check, with the abrupt ethylene treatments only slightly longer than the CIPC check
(126, 0 and 14 mm, respectively). Maximum sprout length in response to temperature
was longer in tubers stored at 13 °C than at 9 °C (14 and 8 mm, respectively).

Early Exposure to Ethylene Gas

Commencing the ethylene exposure early in the storage season (in early October before
suberization was completed) did not darken fry colour as much as the eth check (in
which ethylene exposure began in early December) in both RB and Shp tubers (Table 4).
In RB at the first evaluation date after the start of ethylene exposure, fry colour was 12
vs. 22 ARu darker than the untreated control tubers in the pre-sub abrupt and eth
check treatments, respectively. Recovery to a lighter colour progressed steadily in both
treatments, but the pre-sub abrupt treatment always had lighter fry colour than the eth
check (Table 4). At the end of the storage term, fry colour of RB tubers exposed to
ethylene before suberization ended (pre-sub abrupt) was only 8 ARu darker than the
fry colour of tubers from the CIPC treatment (Table 4a).
In Shp, as in RB, commencing the ethylene exposure earlier in the storage season
had less effect than the eth check on fry colour (Table 4b). Fry colours in the pre-sub
 316

Table 4 Effects of starting ethylene exposure before suberization was completed vs. after suberization and cooling were finished, on potato fry colour and sprouting

Treatment Fry coloura (Agtron % reflectance) Maximum sprout length, Sprout massb,
33–35 weeks (mm) 33–35 weeks (g kg−1)
5 weeksc 8–9 weeks 13–14 weeks 18–19 weeks 23–24 weeks 28–29 weeks 33–35 weeks

a. cv. Russet Burbank, mean of 5 years (1994–1995, 1995–1996, 1997–1998, 1998–1999, 1999–2000)
Untreated control 74.1 ad 73.9 a 74.9 a 77.1 a 77.2 a 77.7 a 76.2 b 316.7e (2.502 a) 45.8e (1.6700 a)
CIPC check –f – 74.0 a 76.6 a 77.2 a 80.2 a 81.4 a 0.0 (0.000 c) 0.0 (0.0000 c)
g
Ethylene
Eth check – – 52.5 c 57.0 c 59.9 c 64.3 c 66.7 d 12.8 (1.141 b) 0.6 (0.1966 b)
Pre-sub abrupt 61.7 b 63.6 b 65.7 b 69.8 b 70.4 b 72.3 b 73.2 c 9.0 (0.998 b) 0.7 (0.2191 b)
SE = 0.916 0.03398h 0.0631h
P= <0.001 <0.001 <0.001

b. cv. Shepody, mean of 2 years (1997–1998, 1998–1999)

7–8c weeks 11–12 weeks 14–15 weeks 17–18 weeks 20–21 weeks 23–24 weeks 26–27 weeks 26–27 weeks 26–27 weeks
Untreated control 87.5 a 86.2 a 85.9 a 83.0 a 83.6 a 84.5 a 83.3 a 121.5 (2.088 a) 22.1 (1.3640 a)
CIPC check – – 83.9 a 82.0 a 82.8 a 78.1 a 82.8 a 0.0 (0.000 c) 0.0 (0.0000 c)
Ethylene
Eth check – – 56.0 c 54.6 c 53.9 c 60.5 c 55.5 c 15.6 (1.219 b) 1.4 (0.3871 b)
Pre-sub abrupt 72.6 b 72.6 b 64.9 b 62.3 b 62.0 b 60.4 c 63.1 b 18.5 (1.289 b) 1.8 (0.4493 b)
SE = 1.946 0.1022 0.03275
P= <0.001 <0.001 <0.001

a
Mean initial fry colours were 76.1±8.4 and 86.4±15.9 Agtron % reflectance units in Russet Burbank (1 week postharvest) and Shepody (3–4 weeks postharvest), respectively
b
Proportion of the initial tuber mass at the start of the trial (mid-October)
c
Weeks after tubers were harvested, treatment × evaluation date interaction
d
For each cv, values within a column followed by the same letter are not significantly different at P≤0.05. To maintain clarity of the fry colour data, differences across evaluation
dates have not been indicated. For these comparisons, three times the SE can be used to closely approximate the least significant difference between means
e
Back-transformed values; log10 means from the statistical analysis are in parentheses
f
Dash indicates no data because this treatment was not yet started
g
Ethylene treatments as described in Table 1
h
SE and P-values for sprouting characteristics pertain to log10 values used in the statistical analysis
Potato Research (2007) 49:303–326
Potato Research (2007) 49:303–326 317

abrupt and eth check were 15 and 30 ARu darker than controls, respectively, at the
first evaluation after the ethylene exposure commenced. However, the magnitude of
darkening in response to ethylene was greater in Shp than in RB, i.e. 15 vs. 12 ARu
darker than controls, respectively, in the pre-sub abrupt treatment, and 30 vs. 22 ARu
darker than controls, respectively, in the eth check. The recovery to a lighter fry
colour observed in RB was not apparent in Shp. Fry colour in the eth check in Shp
was lighter at 23–24 weeks PH than at 14–15 weeks PH, but darkened again at the
final evaluation at 26–27 weeks PH (Table 4b). In the pre-sub abrupt treatment, fry
colour was the same at 7–8 and 11–12 weeks PH, but was 8–11 ARu darker from the
14–15 week PH evaluation until the end of the trials. In contrast, fry colour in the
CIPC check and control tubers remained above 80 ARu throughout the trial (except
CIPC check tubers at 23–24 weeks PH; Table 4b).
Sprout inhibition (sprout mass and maximum sprout length) in the pre-sub abrupt
treatment was equivalent to the eth check in both cultivars (Table 4). Although
sprout mass and maximum length in both ethylene treatments were slightly greater
than in the CIPC check, they were much lower than in the untreated controls.

Gradual Ethylene Exposure using a Concentration Gradient

Compared with untreated control tubers evaluated on the same date, the fry colour
of tubers from the abrupt treatments (eth check, pre-sub abrupt and post-sub abrupt)
was 18, 12 and 15 ARu darker, respectively, at the first evaluation after the start of
each ethylene treatment (Table 5a). In contrast, tuber fry colour at the first
evaluations after the ethylene treatments began in the pre-sub conc, pre-sub conc-
fast and post-sub conc-fast treatments darkened less than their counterparts in the
abrupt treatments [4, 2 and 8 ARu darker than the controls, respectively, at the cor-
responding stage of the ethylene exposure (see Fig. 1)]. Tuber fry colour in the pre-sub
conc treatment was not significantly different from the untreated controls at any
evaluation date (Table 5a). Although the fry colour of tubers in the pre-sub conc-fast
treatment appeared to be slightly darker than tubers in the pre-sub conc treatment at
five of the six evaluation dates, these differences were not significant. Similarly,
although the fry colour of tubers in the post-sub conc-fast treatment was slightly
darker than tubers in the pre-sub conc treatment at all four dates when both were
evaluated, the differences were not significant. In contrast, the fry colour of the eth
check tubers was significantly darker than all other ethylene treatments at 14 weeks
PH and all except the post-sub abrupt treatment at 10 weeks PH (Table 5a).
During subsequent testing of the concentration gradient method using four
cultivars, the eth check significantly darkened tuber fry colour in comparison with
the untreated control and CIPC check samples (Tables 5b,c, and 6), as observed in
previous trials. However, compared with the eth check, the pre-sub conc treatment
reduced the amount of ethylene-induced fry colour darkening in all cultivars,
particularly during the initial weeks of exposure to the gas. This reduction in
darkening was equal to or greater than the improvement attributable to beginning the
ethylene exposure early in the storage season (Tables 5b,c, and 6).
In RB, fry colour of tubers in the eth check was more than 23 ARu darker than
tubers in the CIPC check at 14 weeks PH, the first evaluation after ethylene exposure
began in this treatment (Table 5b). By 34–35 weeks PH, fry colour in the eth check
 318

Table 5 Effects of commencing ethylene exposure by gradually increasing the concentration of ethylene, on potato fry colour and sprouting, cvs Russet Burbank and Shepody

Treatment Fry coloura (Agtron % reflectance) Maximum sprout Sprout massf

b length (mm) (g kg−1)
2 weeks 4 weeks 6 weeks 8 weeks 10 weeks 14 weeks

a. cv. Russet Burbank, 1996–1997 only

Untreated control 71.7 ac 76.3 a 74.3 a 75.6 a 77.9 a 73.4 a
Ethylened
Eth check –e – – – 60.0 d 48.1 d
Pre-sub abrupt 59.3 b 64.0 b 59.0 c 62.7 cd 69.8 abc 62.1 c
Pre-sub conc 68.0 a 70.1 ab 67.6 ab 73.7 ab 75.3 ab 70.8 ab
Pre-sub conc-fast 69.5 a 69.4 ab 63.2 bc 67.8 abc 70.6 abc 64.8 bc
Post-sub abrupt – – 59.5 bc 56.7 d 65.1 cd 63.1 bc
Post-sub conc-fast – – 66.4 abc 65.8 bc 67.3 bc 68.9 abc
SE = 1.946
P= <0.001

b. cv. Russet Burbank, mean of 3 years (1997–1998, 1998–1999, 1999–2000)

5 weeks 9 weeks 14 weeks 19 weeks 24 weeks 29 weeks 34–35 weeks 33–35 weeks 33–35 weeks
Untreated control 72.8 a 72.6 a 72.1 a 74.5 a 73.8 a 74.9 a 72.2 b 270.6g (2.434 a) 46.8 (1.679 a)
CIPC check – – 69.6 ab 71.9 ab 74.8 a 76.9 a 77.7 a 0.0 (0.000 c) 0.0 (0.000 c)
Ethylene
Eth check – – 46.2 d 49.5 d 54.7 c 57.4 c 60.9 d 10.0 (1.040 b) 0.6 (0.195 b)
Pre-sub abrupt 60.8 c 63.0 b 63.4 c 67.5 c 69.3 b 69.6 b 68.5 c 14.8 (1.198 b) 0.8 (0.257 b)
Pre-sub conc 68.2 b 70.5 a 68.2 bc 70.0 bc 70.2 b 71.4 b 71.1 bc 9.3 (1.011 b) 0.4 (0.133 b)
Potato Research (2007) 49:303–326
Table 5 (continued)

Treatment Fry coloura (Agtron % reflectance) Maximum sprout Sprout massf

2 weeksb 4 weeks 6 weeks 8 weeks 10 weeks 14 weeks length (mm) (g kg−1)

SE = 1.172 0.0777h 0.455h

P= <0.001 <0.001 <0.001
c. cv. Shepody, mean of 2 years (1997–1998, 1998–1999)
7–8 weeks 11–12 weeks 14–15 weeks 17–18 weeks 20–21 weeks 23–24 weeks 26–27 weeks 26–27 weeks 26–27 weeks
Untreated control 87.4 a 86.2 a 85.9 a 83.0 a 83.6 a 84.5 a 83.3 a 121.5 (2.088 a) 22.1 (1.3640 a)
CIPC check – – 83.9 a 82.0 a 82.8 a 78.1 b 82.8 a 0.0 (0.000 c) 0.0 (0.000 d)
Ethylene
Eth check – – 56.0 d 54.6 c 53.9 d 60.4 c 55.5 c 15.6 (1.219 b) 1.4 (0.3871 c)
Pre-sub abrupt 72.6 c 72.6 c 64.9 c 62.3 b 62.0 b 60.4 c 63.1 b 18.5 (1.289 b) 1.8 (0.4493 bc)
Pre-sub conc 79.8 b 79.9 b 73.6 b 66.2 b 65.8 b 62.0 b 65.5 b 17.6 (1.270 b) 2.4 (0.5315 b)
SE = 1.927 0.0955h 0.03190h
P= <0.001 <0.001 0.001
Potato Research (2007) 49:303–326

a
Mean initial fry colours were 73.4±11.4, 74.7±9.6 and 86.4±15.9 Agtron % reflectance units in RB section A (1 week postharvest), RB section B (1 week postharvest) and Shp
section C (3–4 weeks postharvest), respectively
b
Weeks after tubers were harvested, treatment × evaluation date interaction
c
For each section of the table, values within a column followed by the same letter are not significantly different at P≤0.05. To maintain clarity of the fry colour data, differences
across evaluation dates have not been indicated. For these comparisons, three times the SE can be used to closely approximate the least significant difference between means
d
Ethylene treatments as described in Table 1
e
Dash indicates no data because this treatment was not yet started
f
Proportion of the initial tuber mass at the start of the trial (mid-October)
g
Back-transformed values; log10 means from the statistical analysis are in parentheses
h
SE and P values for sprouting characteristics only pertain to log10 values used in the statistical analysis
319
 320

Table 6 Effects of ethylene sprout inhibitor treatments on potato fry colour and sprouting, cvs Asterix and Santana

Treatment Fry coloura (Agtron % reflectance) Maximum sprout length (mm) Sprout massb (g kg−1)
6 weeksc 10 weeks 15 weeks 20 weeks 25 weeks 30 weeks 35 weeks 30 weeks 35 weeks 30 weeks 35 weeks

a. cv. Asterix, 1998–1999

Untreated control 86.6 ad 90.0 a 84.9 a 85.3 a 83.8 a 80.9 b 84.4 a 160.3 a 194.0 a 50.0 a 66.8 a
CIPC check –e – 84.9 a 83.6 a 87.7 a 90.6 a 77.1 b 16.0 b 21.8 b 0.7 c 0.7 c
Ethylenef
Eth check – – 40.0 d 47.3 d 53.5 c 57.2 d 42.8 c 40.0 b 41.2 b 14.6 b 31.2 b
Pre-sub abrupt 60.5 c 66.2 c 64.0 c 62.5 c 69.1 b 56.5 d 35.4 d 35.3 b 24.5 b 25.8 b 28.3 b
Pre-sub conc 75.7 b 77.6 b 71.1 b 71.4 b 68.4 b 64.6 c 43.9 c 36.3 b 33.7 b 18.5 b 33.4 b
SE = 2.314 14.60 4.504
P= <0.001 <0.001 <0.001

b. cv. Santana, 1998–1999

Untreated control 89.6 a 91.0 a 89.8 a 90.3 a 91.7 a 91.2 a 90.3 a 142.0 a 184.0 a 30.7 a 52.8 a
CIPC check – – 88.8 a 89.6 a 93.5 a 91.6 a 86.4 ab 9.3 c 13.0 b 0.3 c 0.2 d
Ethylene
Eth check – – 74.0 b 78.7 b 84.7 c 83.0 b 77.3 c 36.0 b 32.7 b 18.5 b 17.8 c
Pre-sub abrupt 82.6 b 85.3 b 86.8 a 86.6 a 86.8 b 87.4 a 88.0 ab 32.5 b 37.5 b 20.5 b 36.3 b
Pre-sub conc 90.2 a 87.9 ab 87.6 a 87.5 a 90.5 ab 87.6 a 84.0 b 33.7 bc 26.0 b 24.1 b 34.1 b
SE = 1.470 8.68 2.115
P= <0.001 <0.001 <0.001

a
Mean initial fry colours were 83.5±4.1 and 90.0±2.2 Agtron % reflectance units in cv. Asterix (2 weeks postharvest) and cv. Santana (2 weeks postharvest), respectively
b
Proportion of the initial tuber mass at the start of the trial
c
Weeks after tubers were harvested, treatment × evaluation date interaction
d
For each cultivar, values within a column followed by the same letter are not significantly different at P≤0.05. To maintain clarity of the fry colour data, differences across
evaluation dates have not been indicated. For these comparisons, three times the SE can be used to closely approximate the least significant difference between means
e
Dash indicates no data because this treatment was not yet started
f
Ethylene treatments as described in Table 1
Potato Research (2007) 49:303–326
Potato Research (2007) 49:303–326 321

had lightened considerably, but it was still 17 ARu darker than the CIPC check at
that date. In contrast, fry colour of tubers in the pre-sub conc treatment was only 1–7
ARu darker than the CIPC check throughout the trials (Table 5b).
In Shp, fry colour of tubers in the pre-sub conc treatment darkened slowly
throughout the trial, and more quickly in the pre-sub abrupt treatment (Table 5c). Fry
colour of tubers in the pre-sub conc treatment was lighter than tubers in the eth
check at all evaluation dates. However, fry colour in the pre-sub conc treatment was
10–17 ARu darker than the CIPC check throughout the trials.
In Ast, fry colour was strongly affected by all ethylene treatments, although the
effect of the pre-sub conc treatment was much less severe than in either the eth check
or the pre-sub abrupt treatment (Table 6a). In the eth check, fry colour was 45 ARu
darker than the CIPC check at 15 weeks PH, but recovered to ca. 34 ARu darker than
the CIPC check at 30 weeks PH. In contrast, fry colour of tubers in the pre-sub conc
treatment was less than 14 ARu darker than the control and CIPC check tubers at 15
and 20 weeks PH, but continued to darken steadily thereafter (Table 6a). At 35 weeks
PH, fry colour in all ethylene treatments in Ast was much darker than at 30 weeks PH.
In San, tuber fry colour in the pre-sub abrupt and pre-sub conc treatments was
lighter than the eth check at all evaluation dates, and was relatively unaffected by
both of the early ethylene treatments in comparison with the CIPC check. In the pre-
sub conc treatment, San fry colour was not significantly different from the CIPC
check at any evaluation date (Table 6b).
Shp, Ast and San all have shorter dormancy than RB, and sprouting began earlier
in these three cultivars than in RB, with or without ethylene sprout inhibitor; many
Ast and San tubers had small sprouts or “pips” visible at many eyes at the start of the
trial in October (data not presented). Nevertheless, sprout elongation was clearly
inhibited by ethylene in all cultivars (Tables 5b,c, and 6). Maximum sprout length in
all four cultivars was much shorter in the ethylene treatments than in the controls.
The early ethylene treatments inhibited sprout growth (both maximum length and
sprout mass) as effectively as the eth check. Unlike RB and Shp, in Ast and San
even the CIPC check tubers sprouted (Table 6).
In Ast and San heavy rosetting at multiple eyes was observed, resulting in
moderate sprout mass at 30 and 35 weeks PH in all ethylene treatments (Table 6).
Despite this fact, sprout mass in the ethylene treatments was still much less than in
the untreated controls in both cultivars.

Gradual Ethylene Exposure using an Exposure-Time Gradient

In both RB and Shp, the fry colour of tubers in the timed method of ethylene
exposure was equivalent to the colour of tubers in the conc method, and was lighter
than the fry colour of tubers in the eth check (Tables 3 and 7). In RB, fry colour of
tubers in the pre-sub timed treatment was as light as tubers in the CIPC check
through 14 weeks PH, inclusive (Table 7). Thereafter, fry colour in the pre-sub timed
treatment was 8–14 ARu darker than tubers in the CIPC check.
In the 1998–1999 temperature trial using Shp, mean tuber fry colour in the mid-
Jan timed treatment at 13 °C was equivalent to the fry colour of tubers in the CIPC
check and the mid-Jan conc treatment at 13 °C, but was lighter than tubers in the eth
check at 9 °C (Table 3a). In the 1999–2000 temperature trial using RB tubers, mean
 322

Table 7 Effects of gradually introducing ethylene by increasing the concentration vs. increasing the duration of exposure, on potato fry colour and sprouting, cv. Russet Burbank
1999–2000

Treatment Fry coloura (Agtron % reflectance) Maximum sprout length (mm) Sprout massb (g kg−1)
5 weeksc 9 weeks 14 weeks 19 weeks 26 weeks 29 weeks 32 weeks 35 weeks 35 weeks 35 weeks

Untreated control 79.0 ad 79.1 a 76.5 a 79.5 a 77.4 ab 80.3 a 78.6 b 74.1 b 269.4e (2.432 a) 42.2e (1.635 a)
CIPC check –f – 76.9 a 76.7 ab 82.2 a 80.8 a 85.1 a 81.6 a 0.0 (0.000 c) 0.0 (0.000 b)
Ethyleneg
Eth check – – 51.8 c 50.1 c 62.4 d 59.0 c 66.2 c 63.1 d 15.2 (1.210 b) 0.2 (0.086 b)
Pre-sub abrupt 68.2 b 68.0 c 68.0 b 72.7 b 73.0 bc 72.3 b 73.1 b 68.1 cd 17.3 (1.262 b) 0.5 (0.173 b)
Pre-sub conc 77.3 a 78.2 ab 73.5 ab 74.9 ab 73.0 bc 73.5 b 76.8 b 73.9 b 7.6 (0.933 b) 0.4 (0.146 b)
Pre-sub timed 78.2 a 73.4 b 74.9 a 73.6 b 68.4 c 71.5 b 76.8 b 70.5 bc 15.5 (1.217 b) 0.4 (0.141 b)
SE = 1.970 0.1356h 0.908h
P= <0.001 0.032 <0.001

a
Mean initial fry colour was 79.6±7.39 Agtron % reflectance units (1 week postharvest)
b
Proportion of the initial tuber mass at the start of the trial (mid-October)
c
Weeks after tubers were harvested
d
Values within a column followed by the same letter are not significantly different at P≤0.05. To maintain clarity of the fry colour data, differences across evaluation dates have
not been indicated. For these comparisons, three times the SE can be used to closely approximate the least significant difference between means
e
Back-transformed values; log10 means from the statistical analysis are in parentheses
f
Dash indicates no data because this treatment was not yet started
g
Ethylene treatments as described in Table 1
h
SE and P-values for sprouting characteristics only pertain to log10 values used in the statistical analysis
Potato Research (2007) 49:303–326
Potato Research (2007) 49:303–326 323

fry colour in the mid-Jan timed treatment at both 9 and 13 °C was similar to the
colour of tubers in the CIPC check at 9 °C (Table 3b). Mean fry colour of tubers in
the eth check appeared darker than the colour of tubers from all other treatments,
although in this trial the differences were not significant. In the 1999–2000 trial
using RB stored at 9 °C only, the fry colour of tubers in the pre-sub timed treatment
was equivalent to the colour of tubers from the pre-sub conc treatment at all
evaluation dates (Table 7). The fry colour of tubers in the pre-sub timed and pre-sub
conc treatments was as light as tubers in the CIPC check through 19 weeks PH,
inclusive. Thereafter, tuber fry colours in the pre-sub timed and pre-sub conc
treatments were 7–14 ARu darker than tubers in the CIPC check (Table 7). At all
evaluation dates, the fry colour of tubers in the pre-sub timed and pre-sub conc
treatments was lighter than the colour of tubers from the eth check.
Sprout inhibition in the timed treatments was equivalent to the eth check in RB
(Tables 3b and 7). Maximum length and sprout mass were slightly greater in the
ethylene treatments than in the CIPC check (Tables 3b and 7); both were much less
than in the untreated controls although the differences in maximum length were not
significant in the temperature trial (Table 3b). A similar trend was observed in Shp,
except that sprout mass was lower in the eth check than in the other ethylene
treatments (Table 3a).

Discussion

The results of the various ethylene application methods consistently showed that the
abrupt exposure of potato tubers to ethylene gas commencing after suberization and
cooling were complete, i.e. the eth check, resulted consistently in darker fry colour,
compared with all other ethylene treatment methods, CIPC checks and untreated
controls. This is consistent with the work of Parkin and Schwobe (1990), Daniels-
Lake et al. (2005) and Prange et al. (1998).
Variability in the degree of response to the ethylene concentration used (i.e. 4 μl l−1)
was evident among cultivars. For example, Shp and especially Ast tubers appeared
to be more sensitive to ethylene than RB with regard to fry colour, but less sensitive
than RB in regard to sprout inhibition (Tables 5 and 6). In contrast, in San tubers
both fry colour and sprouting were less sensitive to ethylene than RB. This agrees
with the findings of Huelin and Barker (1939), Haard (1971), Rylski et al. (1974),
Pritchard and Adam (1994) and Prange et al. (2005b) who reported cultivar
variations in the respiration, sprouting and fry colour responses to ethylene.
Physiological aging of the tubers may have contributed to additional darkening
observed at 35 weeks PH in Ast and San (Table 7). Although tubers of these cultivars
sprouted heavily in all treatments, the extensive rosetting observed in the ethylene
treatments suggests that the tubers were becoming senescent, which likely contributed
to the fry colour darkening. The occurrence of sprouts in the CIPC check tubers in Ast
and San suggests that regardless of the inhibitor employed, sprout control in these two
cultivars may require stronger inhibition than is necessary for RB and Shp.
Increased respiration rates in response to ethylene exposure are commonplace in
plants, particularly in ripening climacteric fruits. However in potato tubers, the
respiration increase peaks rapidly and then declines over the following few days
324 Potato Research (2007) 49:303–326

(Reid and Pratt 1972; Rylski et al. 1974; Daniels-Lake and Prange, unpublished
data), remaining insensitive to additional ethylene exposure during this time (Reid
and Pratt 1972). Since respiration rates reflect changes in tuber sugar concentrations,
which in turn determine fry colour (Habib and Brown 1956; Pritchard and Adam
1994), it was reasonable to hypothesize that interruptions in ethylene exposure could
reduce the negative effect of ethylene sprout inhibitor on fry colour. In the work
reported here, reduced exposure to ethylene by repeated interruptions did indeed
reduce ethylene-induced fry colour darkening but also reduced sprout inhibition. For
example, when ethylene exposure was interrupted for intervals of 3 days out of 4 or 18
h per day throughout the storage season, fry colour was equivalent to the CIPC check,
but sprout growth was greater than in the eth check although still much less than in the
control (Table 2). This methodology could be economically advantageous in the
commercial setting by reducing gas consumption, but the improvement in fry colour
would have to be weighed against the decline in overall quality due to sprouting.
Warmer storage (i.e. 13 °C vs. the standard of 9 °C) reduced ethylene-induced fry
colour darkening, although sprout growth, weight loss and disease were increased in
some cultivars, with or without ethylene (Table 3). These effects are attributable to
an increased rate of respiration, resulting from both the temperature and the direct
effects of the ethylene (Haard 1971; Reid and Pratt 1972; Day et al. 1978; Dwelle and
Stallknecht 1978; Parkin and Schwobe 1990; Pritchard and Adam 1994). The
increased respiration is likely to have consumed much of the additional sugars
resulting from the ethylene exposure which reduced the effect on fry colour, in
addition to favouring sprout growth, weight loss due to both respiration and
transpiration, and the growth of pathogens.
The early introduction of ethylene was effective in maintaining sprout control
comparable to CIPC while reducing fry colour darkening, compared with the eth
check which began at a later date. This is attributable, at least in part, to the effects of
the warm storage temperature during the suberization phase of storage, which
affected respiration, and in turn fry colour, as discussed above.
Gradually introducing the ethylene by either the concentration method or the
timing method similarly reduced the effect of ethylene on fry colour, in comparison with
abrupt exposure to 4 μl l−1 ethylene in the eth check. Low concentrations of ethylene
(e.g. 0.4 μl l−1 or less) do not darken fry colour as much as the 4 μl l−1 concentration
used in the eth check (Daniels-Lake et al. 2005). By starting at a low concentration,
the tubers became acclimated to the presence of ethylene in the storage atmosphere
with only a minimal effect on fry colour. During the later portion of the storage term,
continuous exposure to 4 μl l−1 ethylene effectively inhibited sprout growth.
The timed method of gradually introducing ethylene sprout inhibitor was found to
be equivalent to the concentration method, in terms of both maintaining good fry
colour and inhibiting sprouting. This is despite the fact that the ethylene con-
centration used in the timed method was 4 μl l−1, i.e. the same as used in the abrupt
method. Although the reason for this is not entirely clear, it is likely that when
ethylene was not present in the storage atmosphere, it diffused out of the tubers,
effectively reducing the concentration within the tuber cells.
In terms of tuber quality, there was no clear difference between the two gradual
introduction methods, i.e. increasing the concentration or increasing the daily
exposure time produced similar results. However, increasing the daily exposure time
Potato Research (2007) 49:303–326 325

was technically simpler than increasing the concentration because it did not involve
the analytical challenges of measuring low ethylene concentrations or the difficulties
associated with frequent adjustment of the gas flow-rate into the chambers. This may
be advantageous in a commercial setting.
Combining an early start of ethylene exposure (before the end of suberization)
with either of the gradual introduction methods is judged to be the best ethylene
sprout inhibitor methodology among those tested. This can be expected to provide
good retention of the preferred light fry colour plus effective sprout inhibition.
Within the potato industry, increasing the long-term storage temperature is generally
believed to negatively affect the processing quality of the tubers and favour disease
development. However, the results of this work suggest that further research may be
warranted on early, gradual introduction of ethylene combined with a warmer storage
temperature, to further minimize the impact of ethylene sprout inhibitor on fry colour
while maintaining adequate sprout control.
The observed variability in cultivar responses makes it clear that each cultivar should
be evaluated carefully prior to commercial use of ethylene sprout inhibitor, particularly
if the potatoes are destined for processing. Nevertheless, ethylene holds great promise
for the potato industry to reduce dependence on traditional chemical sprout inhibitors.
The methodologies examined here may serve as tools to help fulfill that promise.

Acknowledgements The authors wish to thank Dr Kenneth McRae for statistical expertise and
assistance, Drs McRae and John Delong for reviewing the manuscript, and Peter Harrison, Chiam Liew,
Robyne Page and Kimberley Hiltz for essential technical contributions. This research was supported in
part by McCain Foods and Cavendish Farms, by material and financial contributions to the work.

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