A

acdittidhs©

K

1

baea(hch

UA

0d

Available online at www.sciencedirect.com

Postharvest Biology and Technology 48 (2008) 199–205

Different responses of goldenberry fruit treated at four maturitystages with the ethylene antagonist 1-methylcyclopropene

Marıa Soledad Gutierrez a, Gustavo D. Trinchero b, Ana Marıa Cerri c,Fernando Vilella c, Gabriel O. Sozzi a,d,∗

a Catedra de Fruticultura, Facultad de Agronomıa, Universidad de Buenos Aires, Avda. San Martın 4453, C1417DSE Buenos Aires, Argentinab Catedra de Bioquımica, Facultad de Agronomıa, Universidad de Buenos Aires, Avda. San Martın 4453, C1417DSE Buenos Aires, Argentina

c Catedra de Produccion Vegetal, Facultad de Agronomıa, Universidad de Buenos Aires, Avda. San Martın 4453,C1417DSE Buenos Aires, Argentina

d CONICET, Argentina

Received 5 August 2007; accepted 4 October 2007

bstract

Goldenberries (Physalis peruviana L.) were harvested at four different maturity stages: immature green (IG), mature green (MG), yellow (Y)nd orange (OR). Fruit were treated or not with 1-methylcyclopropene (1-MCP) at 25 ◦C for 20 h and then kept at 20 ◦C for 8 d. 1-MCP gasoncentrations used were 0, 0.5 or 5 �L L−1. Application of 1-MCP delayed the onset of the ethylene climacteric in IG and MG fruit in a dose-ependent fashion, and transiently decreased ethylene production in Y and OR fruit. 1-MCP application transitorily decreased the respiration raten MG fruit and, to a lesser extent, in IG and Y fruit. In contrast, 1-MCP-treated OR fruit displayed higher respiration rates than control fruithroughout the experimental period. Fruit treated with 5 �L L−1 1-MCP showed for 6 d average CO2 production levels approximately 41% higherhan those of control fruit, with a respiration rate up to 76% higher on day 4. The hue angle was higher in 1-MCP-treated Y goldenberries than

n control fruit for 8 d but differences between MG treated and untreated fruit were recorded only at the end of the experiment. In contrast, noifferences were detected in IG and OR fruit. 1-MCP-treated fruit were not consistently firmer than untreated fruit during storage. Also, 1-MCPad no effect on soluble solids or ascorbic acid contents. 1-MCP application did not prevent decay in OR fruit but reduced its incidence thusuggesting that it may influence pathogen infection and development in ripe goldenberries. 2007 Elsevier B.V. All rights reserved.

yclop

ima

isoc

eywords: Ascorbic acid; Color; Ethylene; Firmness; Goldenberry; 1-Methylc

. Introduction

Goldenberry (Physalis peruviana L.), also called cape goose-erry, is a perennial plant native to tropical South Americand extensively used as part of folk remedies for various dis-ases. The extracts from different parts of the plant shownti-hepatotoxic (Arun and Asha, 2007) and anti-inflammatoryWu et al., 2006) activity as well as anti-proliferative effects on

epatoma cells (Wu et al., 2004). The edible fruits, bearing aomplex volatile profile (Mayorga et al., 2001) and containingigh levels of vitamins A, B, C, �-carotene, phosphorus and

∗ Corresponding author at: Catedra de Fruticultura, Facultad de Agronomıa,niversidad de Buenos Aires, Avda. San Martın 4453, C1417DSE Buenos Aires,rgentina. Tel.: +54 11 4524 8055; fax: +54 11 4514 8739.

E-mail address: gsozzi@agro.uba.ar (G.O. Sozzi).

sedeaggac

925-5214/$ – see front matter © 2007 Elsevier B.V. All rights reserved.oi:10.1016/j.postharvbio.2007.10.003

ropene; Respiration

ron (Hewett, 1993; Sarkar and Chattopadhyay, 1993), are cli-acteric seedy berries (Trinchero et al., 1999) encapsulated in

n inflated, bladder-like calyx called ‘Chinese lantern’.Goldenberry fruit is somewhat unusual as it continues to

ncrease in weight throughout development and ripening whentill attached to the plant. As the fruit color turns from green torange due to chlorophyll breakdown and carotenoid (chiefly �-arotene) accumulation, a progressive softening occurs and theoluble solids/titratable acidity ratio increases linearly (Fischert al., 1997). This fruit shows a conspicuous rise in ethylene pro-uction associated with the climacteric ripening stage (Trincherot al., 1999). Goldenberries vary widely in ethylene productionccording to the cultivar and maturity stage, orange ripe berries

enerally having much higher production rates than immaturereen or mature green berries (Trinchero et al., 1999; Majumdernd Mazumdar, 2002). In climacteric fruits, ethylene closelyoordinates a number of catabolic and anabolic events during

2 logy

rna

oiprSof(afiMotoMg

2

2

oAwwiaslscpt(p

2

toMwdttttbdTm2

wva

atOgplfwt

t1

2

bwePgsT2oef

fASatirwp

MhcmTtr

iflE

00 M.S. Gutierrez et al. / Postharvest Bio

ipening resulting in rapid perishability. Nevertheless, there iso information about the effects of the application of an ethylenection blocker on goldenberry ripening.

1-Methylcyclopropene (1-MCP) is a gaseous ethylene antag-nist – chemically similar to natural occurring substances – thatnactivates the ethylene receptor in different fruits and otherlant tissues for a varying number of days by single exposure toelatively very low dose levels of this cyclic olefin (Sisler anderek, 1997; Sisler, 2006). In the last 5 years, different aspectsf 1-MCP impact on postharvest biology and technology ofruits and other commodities have been extensively reviewedBlankenship and Dole, 2003; Sisler and Serek, 2003; Watkinsnd Miller, 2003; Watkins, 2006; Sozzi and Beaudry, 2007). Therst objective of this work was to examine the potential of 1-CP for control of ripening in goldenberry fruit. The second

bjective was to determine whether goldenberry responsivenesso 1-MCP varies according to the maturity stage. To the bestf our knowledge, this is the first report on the effects of 1-CP application on different ripening and quality indices of

oldenberry fruit.

. Materials and methods

.1. Plant material

Goldenberry fruit were obtained from plants grown in therchard of the Facultad de Agronomıa, Universidad de Buenosires (34◦37′46′′S, 58◦27′46′′W). Seedlings were field-plantedhen they were 15–20 cm tall. Space between plants and rowsas 1 m. Fruits were hand-picked on 15 January 2007, apply-

ng the following selection criteria, as modified from Baumannnd Meier (1993)—stage IG: immature green berry, fresh calyx;tage MG: mature yellowish green berry, greenish to pale yel-ow calyx; stage Y: dark yellow berry, calyx not completely dry;tage OR: orange ripe berry, light brown, dry and paper-likealyx. The inflated calyx was separated from the fruit and dis-osed of. Fruit free from decay and defects were randomizedo provide three 200-fruit experimental units per maturity stageIG, MG, Y or OR). Each 200-fruit unit was placed in a 60-Llastic container.

.1.1. 1-MCP application and fruit storage1-MCP was applied on the day of harvest. The experimen-

al units were kept as untreated controls or treated with 0.5r 5 �L L−1 1-MCP (one experimental unit per treatment). 1-CP was released from 50 mL-capped test vials containingeighed amounts of SmartFreshTM powder (0.14% active ingre-ient; Rohm and Hass, Argentina) by adding warm water (40 ◦C)hrough a septum. Each vial was vortexed and placed in the plas-ic container. The vial was then opened followed by placement ofhe container lid within 10 s. The lid of each container was alsoaped to ensure a tight seal. A commercial sodium hydroxideox (CO2 scrubber) was placed in each container as previously

escribed (Trinchero et al., 2004) to reduce CO2 accumulation.he 1-MCP concentration was verified by means of gas chro-atography, using iso-butylene as standard (Trinchero et al.,

004). Control fruit were kept in 60-L sealed containers having

fi

fs

and Technology 48 (2008) 199–205

ater but no 1-MCP. After 20 h at 25 ◦C, the containers wereented. The experimental units were then stored in air at 20 ◦Cnd 90% RH.

Three 5-fruit groups were picked at random and used forssessment of firmness, color and soluble solids content for eachreatment (0, 0.5, or 5 �L L−1 1-MCP), stage (IG, MG, Y, orR), and ripening period (0, 2, 4, 6, or 8 d). Another three 5-fruitroups per treatment and stage were used to determine ethyleneroduction and respiration rate throughout the experiment. Ethy-ene and respiration assessments were performed daily. Loss ofruit due to decay reduced fruit numbers. Thus, decaying fruitere removed from the containers and recorded at each removal

ime.An additional 180-OR fruit consignment was divided into

hree 60-fruit lots. The lots were treated with 0, 0.5, or 5 �L L−1

-MCP and decay incidence was evaluated after 8 d.

.2. Fruit quality assessment

Ethylene production was determined by placing five golden-erries in a 1.5-L glass container which was in turn tightly sealedith a silicon septum. One milliliter of the head-space gas was

xtracted after 30 min and ethylene was quantified on a Hewlettackard 5890 Series II gas chromatograph (Agilent Technolo-ies, Inc., Santa Clara, CA, USA) fitted with a FID and atainless steel Porapak N column (3.2 mm × 2 m; 80/100 mesh).he injector, oven and detector temperatures were 110, 90, and50 ◦C, respectively. N2 was used as the carrier gas at a flow ratef 0.37 mL s−1 (linear gas velocity = 4.5 cm s−1). Results werexpressed as nanogram of ethylene produced per kilogram ofruit in 1 s.

Gas samples for ethylene and CO2 analyses were collectedrom the same jars. The respiration rate was measured using angilent 4890 gas chromatograph (Agilent Technologies, Inc.,anta Clara, CA, USA) fitted with a TCD and equipped withCarboplotTM (Chrompack) column (0.53 mm × 25 m; 25 �m

hick). Analysis was performed isothermally at 100 ◦C, with thenjector and the detector temperatures held at 80 and 200 ◦C,espectively. He was used as the carrier gas. Linear gas velocityas 68 cm s−1. Results were expressed as microgram of CO2roduced per kilogram of fruit in 1 s.

Hue angle (h◦) values were obtained from the peel with ainolta chromameter (model CR-300; Osaka, Japan) which

as an 8 mm-diameter measuring area. Illuminant C lightingonditions were utilized for the measurements. The chro-ameter was calibrated to a white calibration plate (CR-A43).wo spots on opposite sides of the fruit were measured and

he mean of the two measurements was considered as oneeplicate.

Fruit firmness was measured using an Instron Universal Test-ng Machine (Model 3342, Canton, MA, USA) fitted with aat-plate probe as previously described (Muskovics et al., 2006).ach fruit was compressed 2 mm at a rate of 0.1 mm s−1 and

rmness was determined as previously stated.

Goldenberries were pressed through cheesecloth. The juicerom five goldenberries was pooled and analyzed for solubleolids concentration (SSC) using a hand-held temperature-

M.S. Gutierrez et al. / Postharvest Biology and Technology 48 (2008) 199–205 201

Table 1Indicators of different fruit stages at the beginning of the experiment (mean ± S.E.; n = 3, each replicate containing five fruit)

Stage Mass of one fruit (g) Diameter (mm) Ethylene production (ng kg−1 s−1) Hue angle (h◦)

IG 1.13 ± 0.10 12.90 ± 0.37 3.77 ± 1.32 110.45 ± 1.28MG 1.63 ± 0.10 14.44 ± 0.37 31.25 ± 20.14 103.46 ± 1.97YO

ce

dBHe1cM1XKatar

2

DPITd

3

3

F(tbi

2.47 ± 0.31 16.43 ± 0.72R 3.04 ± 0.21 17.65 ± 0.28

ompensated refractometer (Atago Co., Tokyo, Japan). SSC wasxpressed as a percentage.

Ascorbic acid was extracted and measured as previouslyescribed (Doner and Hicks, 1986), with minor differences.riefly, 2.5 g of fruit was ground with 10 mL of 2% (w/v)Cl in an Omnimixer (Sorvall Inc., Norwalk, CO, USA). The

xtract was immediately centrifuged at 10 000 × g at 4 ◦C for0 min. Aliquots of the supernatant were filtered through aellulose membrane and used for ascorbic acid determination.easurements were performed using a Hewlett Packard Series

100 HPLC (USA) fitted with a reverse-phase column (EclipseDB-C18; 250 mm × 4.6 mm). Methanol:water (30:70; 0.01 M2HPO4, pH 4.30) was used isocratically as the mobile phase

t a flux of 8.3 �L s−1 and the detection was performed spec-rophotometrically. Results were expressed as mass of ascorbiccid per fresh weight mass of fruit, mg kg−1. Three independenteplicates per stage, treatment and date were evaluated.

at1

ig. 1. Ethylene production of control and 1-MCP-treated fruit. Goldenberries were hc) yellow, and (d) orange, exposed or not to 1-MCP and stored at 20 ◦C for 8 d. Numreatment. Values represent the mean ± S.E. Where bars are not shown, the S.E. doesetween T0 and T5 within a single date; letter “b” indicates a significant differencendicates a significant difference between T0 and T5 and between T0.5 and T5 within

199.43 ± 15.24 87.58 ± 1.22346.14 ± 9.72 81.19 ± 1.73

.3. Statistical analysis

Experiments were performed according to a factorial design.ata were analyzed by means of a one-way ANOVA, using theC-SAS software package (SAS Institute Inc., Cary, NC, USA).n the case of a significant F-value, data were subjected to theukey’s test (P < 0.05) for comparison of treatments within eachate.

. Results and discussion

.1. Ethylene production and respiration rate

Goldenberry behaves as a typical climacteric fruit but, asresult of its growth pattern, ethylene increase begins during

he phase of active cell expansion (Table 1; Trinchero et al.,999; Majumder and Mazumdar, 2002). In goldenberry, ethy-

arvested at four different maturity stages: (a) immature green, (b) mature green,bers in the legend indicate the concentration (�L L−1) of 1-MCP used in eachnot exceed the size of the symbol. Letter “a” indicates a significant differencebetween T0 and T0.5, and between T0 and T5 within a single date; letter “c”a single date (P < 0.05).

2 logy

leohit

bpfi

tptcc57hMpits

5

Fstpr

1(suOFdf7ldh(

3

F(tb

02 M.S. Gutierrez et al. / Postharvest Bio

ene biosynthesis can be classified as extremely high (Trincherot al., 1999). Nevertheless, ethylene biosynthesis may dependn the harvest time: goldenberry ethylene production reachedigher levels when harvested in summer (this experiment) thann autumn (Trinchero et al., 1999). This may be attributable tohe variation in ambient temperature conditions during ripening.

At 20 ◦C and 90% RH, untreated preclimacteric IG golden-erries had a lag time of approximately 2 d prior to the ethyleneroduction increase (Fig. 1a). Exposure to 0.5 �L L−1 1-MCPor 20 h lengthened the lag phase by 1 d while 5 �L L−1 1-MCPnhibited the ethylene rise throughout the experimental period.

Untreated MG fruit showed a lag phase of only 1 d prior tohe ethylene production increase (Fig. 1b). MG fruit ethyleneroduction appeared to have a more typical climacteric pat-ern in comparison with that of IG fruit. The onset of ethylenelimacteric was progressively delayed with increasing 1-MCPoncentrations. In the MG stage, fruit treated with 0.5 and�L L−1 1-MCP showed different ethylene productions on daysand 8 (Fig. 1b). On day 3, ethylene synthesis was 15.7-fold

igher in control fruit than in fruit treated with 0.5 �L L−1 1-CP. Fruit treated with 1-MCP did not show a peak in ethylene

roduction but only a gradual rise towards the end of the exper-mental period. In fact, not only did 1-MCP treatment delay

he ethylene climacteric, but also altered the rates of ethyleneynthesis.

Differences between control fruit and fruit treated with�L L−1 1-MCP were significant on days 5–8 (IG stage,

MbCo

ig. 2. Respiration rate of control and 1-MCP-treated fruit. Goldenberries were harvc) yellow, and (d) orange, exposed or not to 1-MCP and stored at 20 ◦C for 8 d. Numreatment. Values represent the mean ± S.E. Where bars are not shown, the S.E. doesetween T0 and T5 within a single date; letter “b” indicates a significant difference b

and Technology 48 (2008) 199–205

ig. 1a), 2–8 (MG stage, Fig. 1b), but only on days 2 and 3 (Ytage, Fig. 1c) and 1–3 (OR stage, Fig. 1d) (P < 0.05). Applica-ion of 5 �L L−1 1-MCP yielded transient decreases in ethyleneroduction of ∼29% and ∼35% in Y and OR goldenberries,espectively (Fig. 1c–d), but this effect only lasted 2–3 d.

Although CO2 production was not consistently affected by-MCP at the IG stage throughout the experimental periodFig. 2a), 1-MCP-treated fruit harvested at the IG, MG and Ytages showed an average respiration rate lower than that inntreated fruit (Fig. 2a–c). In contrast, the respiration rate ofR fruit increased (P < 0.05) in all 1-MCP treatments (Fig. 2d).ruit treated with 5 �L L−1 1-MCP showed average CO2 pro-uction levels approximately 41% higher than those of controlruit from day 1 through day 6, with a respiration rate up to6% higher on day 4. This effect was not sustained during theast 2 d of the experiment. Most climacteric fruit species displayecreased respiration rates upon application of 1-MCP, althoughigher respiration rates were detected in 1-MCP-treated ripe figsSozzi et al., 2005).

.2. Color, firmness, and decay

No differences in h◦ were detected between control and 1-

CP-treated fruit in IG and OR goldenberries (Fig. 3a and d)

ut 1-MCP delayed h◦ change in MG and Y fruit (Fig. 3b and c).ontrol MG fruit displayed a yellowish color at day 4 and a fullrange color at day 8. In contrast, MG goldenberries treated with

ested at four different maturity stages: (a) immature green, (b) mature green,bers in the legend indicate the concentration (�L L−1) of 1-MCP used in eachnot exceed the size of the symbol. Letter “a” indicates a significant difference

etween T0 and T0.5, and between T0 and T5 within a single date (P < 0.05).

M.S. Gutierrez et al. / Postharvest Biology and Technology 48 (2008) 199–205 203

Fig. 3. Hue angle of control and 1-MCP-treated fruit. Goldenberries were harvested at four different maturity stages: (a) immature green, (b) mature green, (c) yellow,and (d) orange, exposed or not to 1-MCP and stored at 20 ◦C for 8 d. Numbers in the legend indicate the concentration (�L L−1) of 1-MCP used in each treatment.Values represent the mean ± S.E. Letter “a” indicates a significant difference between T0 and T5 within a single date; letter “b” indicates a significant differencebetween T0 and T0.5, and between T0 and T5 within a single date (P < 0.05).

Fig. 4. Firmness of control and 1-MCP-treated fruit. Goldenberries were harvested at four different maturity stages: (a) immature green, (b) mature green, (c) yellow,and (d) orange, exposed or not to 1-MCP and stored at 20 ◦C for 8 d. Numbers in the legend indicate the concentration (�L L−1) of 1-MCP used in each treatment.Values represent the mean ± S.E. In all cases, differences between treatments within a single date were not significant (P < 0.05).

2 logy

5efa5Od

bfalaogit

s

Ftmocmcd1

MnMsfdo0aaip

3

04 M.S. Gutierrez et al. / Postharvest Bio

�L L−1 1-MCP had not yet reached the Y stage at the end of thexperiment but showed a h◦ similar to that observed in controlruit on day 2 (Fig. 3b). Also, Y control fruit reached the colorssociated with the OR stage after 4 d while fruit treated with�L L−1 1-MCP showed the same color after approximately 7 d.R control fruit displayed further color development towards aark orange stage (over-ripe).

Treatment with 5 �L L−1 1-MCP tended to delay golden-erry softening in all stages though differences with controlruit were not significant, probably due to a relatively high vari-bility (Fig. 4). Application of 5 �L L−1 1-MCP brought aboutittle effect in tomato, another solanaceous species (Tassoni etl., 2006). Given the physiological variability between stages,ptimum concentrations for 1-MCP treatment are likely to bereater for some maturity stages than for others. Further stud-

es are required to determine if concentrations higher than thoseested in this experiment will maintain firmness.

Decay increased in treated and untreated OR fruit duringtorage but was consistently lower in 0.5 and 5 �L L−1 1-

ig. 5. Soluble solid concentration (a) and ascorbic acid content (b) in con-rol and 1-MCP-treated fruit. Goldenberries were harvested at four different

aturity stages (immature green, mature green, yellow, and orange), exposedr not to 1-MCP and stored at 20 ◦C for 8 d. Numbers in the legend indicate theoncentration (�L L−1) of 1-MCP used in each treatment. Values represent theean ± S.E. Values at day 0 were the same for all treatments. Soluble solids

ontent increased significantly in all treatments between days 0 and 8, but noifferences were detected between treatments within a single date (P < 0.05).-MCP did not affect ascorbic acid content significantly in any stage.

tcAp1tCadtn

tbaACgsa

4

gl(boregacdsrab

f

and Technology 48 (2008) 199–205

CP-treated fruit (data not shown). Even when data showedo statistical differences in firmness between untreated and 1-CP-treated fruit, a greater percentage of the untreated fruit

howed unacceptable decay. In another experiment, three 60-ruit lots were treated with 0, 0.5, or 5 �L L−1 1-MCP andecay incidence was evaluated after 8 d. By day 8, incidencef decay was 35 ± 2%, 15 ± 1% and 15 ± 1% in fruit from 0,.5 and 5 �L L−1 1-MCP treatments, respectively. 1-MCP mayct at different levels within the intricate physiology of the hostnd the host–pathogen complex, decreasing fruit susceptibil-ty through its effect on fruit natural barriers or impacting onathogen metabolic functions (Sozzi and Beaudry, 2007).

.3. Soluble solids content and ascorbic acid

SSC and ascorbic acid concentrations were similar amongreatments within a single date (data not shown). To detecthanges along time, data were also analyzed by a two-wayNOVA. In every maturity stage, SSC showed an increasingattern towards the end of the experimental period (Fig. 5a).-MCP did not prevent SSC increase regardless the concentra-ion of 1-MCP and the maturity stage of the fruit. AlthoughO2 was overproduced in OR 1-MCP-treated fruit (Fig. 2d) andn effect on carbohydrate metabolism could be expected, noifferences in SSC were detected between control and 1-MCP-reated fruit thus suggesting that the respiratory substrates wereon-limiting.

In goldenberries, ascorbic acid content increased from the IGo the OR stage (Fig. 5b). Once the fruit was harvested, ascor-ic acid remained stable throughout the experiment and 1-MCPpplication did not affect its content significantly in any stage.scorbic acid is the chief biologically active form of vitaminand also one of the most abundant antioxidants present in

oldenberries: between 45% and 55% of the total free radical-cavenging activity of goldenberry can be ascribed to ascorbiccid (Trinchero, unpublished data).

. Conclusion

We conclude that (1) climacteric ethylene increase in MGoldenberries is arrested by 1-MCP thus suggesting that ethy-ene production during goldenberry ripening is autocatalytic;2) some of the biochemical changes associated with golden-erry ripening (i.e. color change) are at least partially dependentn functioning ethylene receptors; (3) the effect of ethyleneeceptor blockage strongly depends on the fruit stage: OR gold-nberries show less responsiveness to 1-MCP than MG or Yoldenberries; (4) goldenberry is not highly sensitive to rel-tively low (0.5 �L L−1) doses of 1-MCP and further dosealibration may be needed in future experiments; moreover, theoses utilized could be partially responsible for the relativelylight postponement of fruit softening; (5) 1-MCP-treated fruitesumed ripening, and some attributes related to flavor (SSC)

nd nutritional quality (ascorbic acid content) were not affectedy 1-MCP application.

The efficacy of 1-MCP doses up to 5 �L L−1 in delayingurther softening of MG, Y or OR goldenberries was not consis-

Biolo

tBfpoifb

A

Nc

R

A

B

B

D

F

H

M

M

M

S

S

S

S

S

S

T

T

T

W

W

W

M.S. Gutierrez et al. / Postharvest

ent enough to recommend commercial application on that basis.esides, MG goldenberries attached to the plant increase their

resh weight by 86% towards the OR stage and, consequently,remature detachment of fruit does not seem to be a practicalption. Nevertheless, 1-MCP treatment may influence pathogennfection and development in OR goldenberries, extending theully ripe stage before fruit tissues undergo senescence andecome an easy target for pathogen attack.

cknowledgements

G.O.S. wishes to thank CONICET, UBA and the Agenciaacional de Promocion Cientıfica y Tecnologica for their finan-

ial support.

eferences

run, M., Asha, V.V., 2007. Preliminary studies on antihepatotoxic effect ofPhysalis peruviana Linn. (Solanaceae) against carbon tetrachloride inducedacute liver injury in rats. J. Ethnopharmacol. 111, 110–114.

aumann, T.W., Meier, C.M., 1993. Chemical defense by withanolides dur-ing fruit development in Physalis peruviana. Phytochemistry 33, 317–321.

lankenship, S.M., Dole, J.M., 2003. 1-Methylcyclopropene: a review. Posthar-vest Biol. Technol. 28, 1–25.

oner, L.W., Hicks, K.B., 1986. High-performance liquid chromatographicseparation of ascorbic acid, erythorbic acid, dehydroascorbic acid, dehy-droerythorbic acid, diketogulonic acid, and diketogluconic acid. MethodsEnzymol. 122, 3–10.

ischer, G., Ludders, P., Gallo, F., 1997. Qualitatsveranderungen der kapstachel-beere wahrend der fruchtreifung. Erwerbs-Obstbau 39, 153–156.

ewett, E.W., 1993. New horticultural crops in New Zealand. In: Janick, J.,Simon, J.E. (Eds.), New Crops. John Wiley & Sons, Inc., New York, pp.57–64.

ajumder, K., Mazumdar, B.C., 2002. Changes of pectic substances in devel-

oping fruits of cape-gooseberry (Physalis peruviana L.) in relation to theenzyme activity and evolution of ethylene. Sci. Hort. 96, 91–101.

ayorga, H., Knapp, H., Winterhalter, P., Duque, C., 2001. Glycosidically boundflavor compounds of cape gooseberry (Physalis peruviana L.). J. Agric. FoodChem. 49, 1904–1908.

W

gy and Technology 48 (2008) 199–205 205

uskovics, G., Felfoldi, J., Kovacs, E., Perlaki, R., Kallay, T., 2006. Changes inphysical properties during fruit ripening of Hungarian sweet cherry (Prunusavium L.) cultivars. Postharvest Biol. Technol. 40, 56–63.

arkar, T.K., Chattopadhyay, T.K., 1993. Correlation studies on cape gooseberry(Physalis peruviana L.). Ann. Agric. Res. 14, 211–214.

isler, E.C., 2006. The discovery and development of compounds counteractingethylene at the receptor level. Biotechnol. Adv. 24, 357–367.

isler, E., Serek, M., 1997. Inhibition of ethylene responses in plants at a receptorlevel: recent developments. Physiol. Plant 100, 577–582.

isler, E.C., Serek, M., 2003. Compounds interacting with the ethylene receptorin plants. Plant Biol. 5, 473–480.

ozzi, G.O., Beaudry, R.M., 2007. Current perspectives on the use of 1-methylcyclopropene in tree fruit crops: an international survey. StewartPostharvest Rev. 3 (2), 1–16 (16).

ozzi, G.O., Abrajan-Villasenor, M., Trinchero, G.D., Fraschina, A.A., 2005.Postharvest response of ‘Brown Turkey’ figs (Ficus carica L.) to the inhibi-tion of ethylene perception. J. Sci. Food Agric. 85, 2503–2508.

assoni, A., Watkins, C.B., Davies, P.J., 2006. Inhibition of the ethylene responseby 1-MCP in tomato suggests that polyamines are not involved in delayingripening, but may moderate the rate of ripening or over-ripening. J. Exp.Bot. 57, 3313–3325.

rinchero, G.D., Sozzi, G.O., Cerri, A.M., Vilella, F., Fraschina, A.A., 1999.Ripening-related changes in ethylene production, respiration rate and cell-wall enzyme activity in goldenberry (Physalis peruviana L.), a solanaceousspecies. Postharvest Biol. Technol. 16, 139–145.

rinchero, G.D., Sozzi, G.O., Covatta, F., Fraschina, A.A., 2004. Inhibitionof ethylene action by 1-methylcyclopropene extends postharvest life of“Bartlett” pears. Postharvest Biol. Technol. 32, 193–204.

atkins, C.B., 2006. The use of 1-methylcyclopropene (1-MCP) on fruits andvegetables. Biotechnol. Adv. 24, 389–409.

atkins, C.B., Miller, W.B., 2003. Implications of 1-methylcyclopropene reg-istration for use on horticultural products. In: Vendrell, M., Klee, H.,Pech, J.C., Romojaro, F. (Eds.), Biology and Technology of the Plant Hor-mone Ethylene, III. IOS Press, Amsterdam, The Netherlands, pp. 385–390.

u, S.J., Ng, L.T., Lin, D.L., Wang, S.S., Lin, C.C., 2004. Physalis peruvianaextract induces apoptosis in human Hep G2 cells through CD95/CD95Lsystem and mitochondrial signalling transduction pathway. Cancer Lett. 215,

199–208.

u, S.J., Tsai, J.Y., Chang, S.P., Lin, D.L., Wang, S.S., Huang, S.N., Ng, L.T.,2006. Supercritical carbon dioxide extract exhibits enhanced antioxidant andanti-inflammatory activities of Physalis peruviana. J. Ethnopharmacol. 108,407–413.