Alleviating sunburn injury in apple fruit using natural and fertilizer forms of S-abscisic acid and its underlying mechanism

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1 WFL Pulisher Siene nd Tehnology Meri-Rstilntie, FI-9 Helsinki, Finlnd e-mil: Journl of Food, griulture & Environment Vol. () : lleviting sunurn injury in pple fruit using nturl nd fertilizer forms of S-sisi id nd its underlying mehnism Kusol Imsu,*, Yoshiko Sekozw, Sumiko Sugy, Hiroshi Gemm nd Ysuo Kmuro Lortory of Pomology, Grdute Shool of Life nd Environmentl Sienes,University of Tsuku, Tsuku -, Jpn. Thilnd Institute of Sientifi nd Tehnologil Reserh, Pthumthni, Thilnd. L Plnning o., Ltd., ihi 49-94, Jpn.*e-mil: kusolimsu@yhoo.om Reeived Jnury 9, epted Mrh 9. strt Twelve-yer-old pple (Mlus pumil Mill.) trees ultivr Tsugru nd grown t the University of Tsuku, Jpn, were sujeted to high tempertures nd high light intensity during summer, fter tretment with ppm nturl S-sisi id or S- [-(-hydroxy-,6,6- trimethyl-4-oxo--ylohexen--yl)--methyl-,4-pentdienoi id] or with 4 nd ppm fertilizer [MIYOI, new fertilizer formultion ontining.,.,.9,. nd. %.i.(w/w) of K, P, Mg, nd Mn, respetively, nd. %.i. (w/w) S-] dys efore hrvesting fruit. ll tretments effetively suppressed the development of sunurn injury mnifested s nerosis in Tsugru nd rowning in s ompred to untreted fruit (ontrol). Totl ntioxidnt pity s well s sori id, totl phenols, hlorophyll nd nd totl nthoynin ontents were higher while mlondildehyde ontent, produt of lipid peroxidtion, ws lower in -treted fruit thn in the ontrol. These results suggest tht inresed ntioxidnt levels to inhiit sunurn injury, disorder resulting from oxidtive ell dmge used y high temperture nd solr rdition. Key words: ntioxidnts, nthoynin, hlorophyll, mlondildehyde, fertilizer omposition, lipid peroxidtion, Mlus pumil, rowning, nerosis, fruit surfe temperture. Introdution pple (Mlus pumil Mill.), temperte fruit, is sensitive to high tempertures nd high levels of solr irrdition, whih previl during summer nd n use sunurn of the fruit peel. s reported previously for ultivrs Grnny Smith, Fuji nd Royl Gl, sunurn is serious prolem in mny res of the world where pples re grown in the presene of high levels of solr irrdition nd high temperture. Glenn et l. suggested tht the use of this dmge ws exess het nd rdition (minly UV-). Sunurn symptoms pper in two min forms. The first type of symptom ppers s nerosis on the fruit surfe. High fruit surfe tempertures (FSTs) of up to ± result in het-indued ell deth, using drk-rown or lk spots severl millimeters in thikness. Symptom development does not require diret exposure to sunlight s het soure. The seond type of symptom, whih does not involve ell deth, is hrterized y yellowing or rowning of the skin. It ours t lower FSTs (46 to 49 ) thn those required for neroti symptom development 4,. Some mesures frequently used to ontrol sunurn for wide vriety of rops, inluding pples, re the use of evportive ooling, kolinsed prtile films (e.g. Surround WP) nd shde nets 6. However, these ontrol mesures hve limittions. In the se of Surround WP film, the umultion of nthoynins hs sometimes een redued, suggesting tht the inresed refletne used y the film uses light limittion,. lthough evportive ooling works well to redue het dmge, it hs no effet on dmge used y solr rdition nd my e diffiult to use in res with wter supply limittion. Shde nets lso work well, ut re expensive, diffiult to implement over lrge res or with lrge trees nd ould redue fruit olor. We previously found tht pplition of the nturl form of sisi id or S- [- (-hydroxy-,6,6-trimethyl-4-oxo--ylohexen--yl)--methyl-,4- pentdienoi id] nd fertilizer [fertilizer formultion ontining.,.,.9,.,. nd.%.i. (w/w) of K, P, Mg,, Mn nd S-, respetively] efore the plnts were sujeted to het nd light stress ould prevent sunurn. This study imed to eluidte the physiologil nd iohemil mehnisms y whih inhiits sunurn injury, therey improving understnding of sunurn protetion nd filitting the development of improved griulturl prties. Mterils nd Methods Plnt mterils: Twelve-yer-old pple trees, ultivr Tsugru nd, grown under field onditions t the griulturl Reserh enter, University of Tsuku, in Jpn s Irki Prefeture (6 N, 4 E), were used. pplitions: oth the fruits nd leves of the trees were spryed with one of the following tretments on July for Tsugru nd 9 ugust for, orresponding to dys efore their respetive hrvesting times; ppm S-, 4 ppm fertilizer, ppm fertilizer nd wter s ontrol. To protet the fruits from insets nd diseses they were overed 446 Journl of Food, griulture & Environment, Vol. (), pril 9

2 with white pper gs immeditely fter or wter spry. The fruits were hrvested t ommeril mturity, nd the peel nd pulp were seprted, frozen in liquid nitrogen nd stored t until hemil nlysis. Methods of hemil nlysis: Mlondildehyde (M), produt of lipid peroxidtion, ws extrted from the peel nd quntified. Frozen peel tissues ( g) were homogenized with ooled pestle in mortr ontining ml of ie-old.% trihloroeti id. The superntnt ws thoroughly mixed with 4 ml of % trihloroeti id ontining.% -thiorituri id. The mixture ws inuted t 9 for min nd then quikly ooled in n ie-th. Totl ntioxidnt pity ws mesured y using,-diphenyl- -pirylhydrzyl (PPH). The rdil-svenging tivity of the extrts from pple peel ws mesured ording to the proedure desried y Zheng nd Wng 4. Totl phenolis ontents in the wter-, ethnol- nd ethyl ette solule extrts of peel tissues were determined y using Folin iolteu regent following the method of Slinkrd nd Singleton nd lulted s mg of glli id equivlent, expressed s mg glli id/g fresh weight of the plnt mteril. For nlysis of totl sori id ontent peel smple ( g) ws mixed with ml of oxli id (%), homogenized for min, filtered, nd ml filtrte dded with mg of polyvinylpolypyrrolidone to remove phenoli ompounds. sori id ontent ws nlyzed y the method of Pntelidis et l.. Totl nthoynin ontent ws determined y the method desried y Pdmvti et l. 9 with slight modifitions. Peel smple ( g) ws extrted with ml of methnol idified with % Hl for 4 h t room temperture. For nlysis of totl hlorophyll, hlorophyll nd hlorophyll ontents, frozen peel smple ( g) ws mixed with ml of N,Ndimethylformmide nd stored for 4 h t 4 in the drk. Spetrophotometri mesurements were mde in triplite for eh smple with Jso V- spetrophotometer (Jso, Tokyo, Jpn). The sorne ws reorded for lipid peroxidtion ( nd 6 nm), totl ntioxidnt pity (6 nm), totl phenoli ontent (6 nm), sori id (4 nm), totl nthoynin ontent ( nm), totl hlorophyll ( nm), hlorophyll (64 nm) nd hlorophyll (664 nm). Sttistil nlysis: Results were nlyzed using SS (Version 9, SS Institute, ry, N, US) for one-wy NOV to test the signifine (p <.) of men differenes. For multiple-men omprisons, Tukey s HS test ws used for M, phenoli, sori id, totl nthoynin, hlorophyll, hlorophyll, nd totl hlorophyll ontents, nd hlorophyll / rtio, nd Sheffé s test for perentge nerosis nd perentge rdil svenging tivity. Results nd isussion Sunurn inidene: ll tretments signifintly redued nerosis nd rowning symptoms of sunurn in oth pple ultivrs (Fig. ). Summer tempertures in ugust nd the diurnl hnges in ir nd fruit temperture on ugust re shown in Fig.. In oth yers, ir tempertures exeeded for ll or most of the study period. Similrly high FSTs were Perentge of sunurn 6 4 ontrol S- ppm Fertilizer 4 ppm Fertilizer ppm Figure. Inhiition of sunurn y S- nd -fertilizer pplition, s expressed y the perentge of sunurned fruit in Tsugru pple. Vlues represent mens ± SE (n = 4). rs (mens ± SE, n = 4) with different letters differ signifintly (Sheffé s test, p <.). ir temperture ( ) ir temperture ( ) Highest temp oserved t the top nd middle prts of the pples, with or without the protetive gs, with vlues rnging from 44 to t the top of the pples nd from to t the middle prt. High FST nd intense sunlight pled oth gged nd re fruits t risk of sunurn. ifferenes in FSTs etween the top nd middle prts of the fruits might reflet differenes in the exposure of the fruits to solr rdition. In, up to 4.% of Tsugru fruits were injured, resulting in nerosis. Het nd solr rdition (minly UV-) re oth known to indue sunurn of pple fruit. Sunurn July, July, ugust, ugust, Septemer, Septemer, (dy (dy of of month) month) g top g mid re top re mid ir-temp Lowest temp :4 : : : : : :4 : : : : : :4 July, ugust, Septemer, (dy of month) Figure. () ir tempertures in ugust ; nd () the ir temperture nd fruit surfe temperture (FST) of gged nd re fruits in Tsugru pple t Tsuku, Jpn, on ugust. Journl of Food, griulture & Environment, Vol. (), pril 9 44

3 generlly ours when the fruit s skin temperture exeeds 4 ; this oservtion is in greement with our results. Mny studies of sunurn in pple hve reported sunurn inidenes rnging from to %,. M ontent: oth ultivrs hd redued M ontents in response to tretments, exept Tsugru fruit treted with S- (Fig. ). M, produt of lipid peroxidtion (or oxidtive degrdtion of lipids), is n inditor of the degree of plnt oxidtive stress. The -indued redution of M ontent oserved in this study indites redued oxidtive stress, possile protetive mehnism ginst high temperture-indued ell dmge due to lipid peroxidtion whih eomes visile s sunurn symptoms. High levels of M equivlent in response to stress hve een reported in mny previous studies nd oinided with high levels of polyunsturted ftty ids in the plnt ells 6. Free ftty ids umulte in plnt memrnes fter exposure to environmentl stresses suh s high or low temperture. Free-rdil retions tht tlyze the seletive degrdtion of phospholipids hve een oserved fter periods of environmentl stress or during plnt senesene. Free rdils from the lipids in ell memrnes use oxidtive dmge to these lipids, resulting in ell dmge. M ontent (nmol-g - fresh wt) M ontent (nmol-g - fresh wt) ntioxidnt pity: signifintly improved the totl ntioxidnt pity or rdil-svenging tivity ompred with Tsugru Figure. Mlondildehyde (M) ontent of fruit peel of Tsugru nd pples hrvested dys fter S- nd -fertilizer pplition. rs (mens ± SE, n = 4) with different letters differ signifintly (Tukey s HS test, p <.). tht of the ontrol in oth ultivrs (Fig. 4). This result suggests tht promoted iosynthesis of ntioxidnts nd/or delyed their deomposition. It hs een reported tht plnts protet their ellulr nd suellulr systems ginst the ytotoxi effets of oxygen rdils y using ntioxidnt enzymes, suh s superoxide dismutse, sorte peroxidse, glutthione redutse nd tlse, nd metolites suh s glutthione, sori id, α- toopherol, rotenoids nd nthoynins. ntioxidnts n lso slow or prevent oxidtion of imoleulr from eing oxidized nd thus the ntioxidnt hs fulfilled its funtion. This protetion n e explined y the pity of ntioxidnts to svenge free rdils, whih re responsile for oxidtive dmge to lipids, proteins nd nulei ids 9. In the present study, the high levels of totl ntioxidnt pity due to tretment my hve ontriuted to the redution sunurn injury. M ontent (nmol-g - fresh wt) M ontent (nmol-g - fresh wt) Totl phenoli ontent: In Tsugru, totl phenoli ontent ws highest with ppm fertilizer, followed y 4 ppm fertilizer nd ppm S-, nd ws lowest in the ontrol (Fig. ). showed different pttern, with only ppm fertilizer nd -ppm S- tretments produing signifintly Tsugru Figure 4. Totl ntioxidnt pity, expressed s rdil svenging perentge, in the fruit peel of Tsugru nd pples hrvested dys fter S- nd -fertilizer pplition. rs (mens ± SE, n = 4) with different letters differ signifintly (Sheffé s test, p <.). 44 Journl of Food, griulture & Environment, Vol. (), pril 9

4 Phenoli ontent (mg-g - fresh wt) Phenoli ontent (mg-g - fresh wt) d d Tsugru Figure. Totl phenoli ontent of fruit peel of Tsugru nd pples hrvested dys fter S- nd -fertilizer pplition. Vlues re in mg glli id equivlent. rs (mens ± SE, n = 4) with different letters differ signifintly (Tukey s HS test, p <.). sori id (mg/g fresh wt) sori id (mg/g fresh wt) Tsugru Figure 6. sori id ontent of fruit peel of Tsugru nd pples hrvested dys fter S- nd -fertilizer pplition. rs (mens ± SE, n = 4) with different letters differ signifintly (Tukey s HS test, p <.). higher phenoli ontents thn the ontrol (Fig. ). Phenolis re seondry plnt metolites tht ontriute to the olor (pperne), tste (itterness) nd texture (stringeny). olor, whih reflets nthoynin ontent, is lso n importnt qulity inditor for fruit. ording to ohen et l. 4, high temperture redues phenoli ontent in grpes. Our results suggest tht supports the umultion of phenoli ompounds in pple peel. The presene of phenoli ompounds in plnts is importnt for norml growth nd development, s these ompounds defend plnt tissues ginst infetion nd injury nd my ply importnt roles in the plnt s oxidtive stility nd in the present study, suppressing or lleviting sunurn dmge. sori id ontent: sori id ontent ws highest in Tsugru treted with ppm fertilizer nd in treted with ppm S- (Fig. 6). In previous study in tomto, sori id ontent deresed in response to high tempertures. These findings imply tht high tempertures use dmge y reduing sori id level nd onsequently ntioxidnt tivity whih ould e prtly reversed y. Totl nthoynin ontent: nthoynin ontent vried widely with tretment (Fig. ). It ws signifintly higher with fertilizer thn with S- in oth ultivrs. nthoynins ply signifint role in the development of fruit olor. In previous study, nthoynin umultion deresed when the proportion of sunurned Fuji pples inresed fter exposure to high tempertures. Het tretment of strwerries ws lso found to inrese the rte of dey of the fruit pulp nd delyed fruit softening, nthoynin umultion nd olor development 9. nthoynin-relted pigments serve s sreen ginst UV light nd ply role in proteting the plnt s N ginst dmge y sunlight, 4,. The -indued inrese in nthoynin ontent s otined in the present study suggests tht fvored nthoynin iosynthesis whih ould hve further ontriuted to its protetive effet ginst sunurn dmge. hlorophyll ontents: hlorophyll ontent inresed in ll treted fruit (Figs. nd 9). The hlorophyll frtions were primrily hlorophyll nd. hlorophyll ws signifintly higher in ll tretments thn the ontrol in Tsugru ut this ws not the se in. hlorophyll ws lso signifintly higher in ll tretments thn the ontrol for oth ultivrs ut differenes mong the three tretments were not signifint. Totl hlorophyll levels were similrly higher in -treted fruit in Tsugru while in only ppm S- used signifint inrese reltive to the ontrol. hlorophyll nd rtio ws signifintly lower in ll tretments thn the ontrol in oth ultivrs. This prtiulr result indites tht inresed hlorophyll ontent my e prtly responsile for the redued sunurn symptoms in the treted fruit. Felietti nd Shrder reported tht in Fuji pple the levels of hlorophyll Journl of Food, griulture & Environment, Vol. (), pril 9 449

5 nthoynin (mg-g - fresh wt) nthoynin (mg-g - fresh wt) 4 4 Tsugru nd oth deresed when the proportion of sunurned fruits inresed. Most plnts ontin two to three times more hlorophyll thn hlorophyll. hl is synthesized y hlorophyllide oxygense (O). O is Rieske-type monooxygense tht onverts hlorophyll into hlorophyll in two-step oxygention retion. UV- exposure deresed hlorophyll levels nd redued the hlorophyll to rtio in n sortedefiient mutnt of ridopsis thlin (vt) ompred with those in the wild-type (WT) plnts 6,. Under supplementry UV- tretment, -week-old vt mutnts showed higher levels of lipid peroxidtion nd H O prodution thn WT plnts 6. hlorophyll ontent in the perirp of tomto fruits deresed during -dy hot ir tretments t 6 to 4. We found higher hlorophyll levels when ws pplied, inditing tht ould either preserve hlorophyll in the fruit peel or promote its prodution, espeilly in the se of hl, whih my ply role in the plnt s defense system. Figure. nthoynin ontent of fruit peel of Tsugru nd pples hrvested dys fter S- nd -fertilizer pplition. rs (mens ± SE, n = 4) with different letters differ signifintly (Tukey s HS test, p <.). lorophyll (mg/g fresh wt) hlorophyll (mg/g fresh wt). Totl lorophyll (mg/g fresh wt) Totl lorophyll (mg/g fresh wt) lorphyll / hlorophyll /... Figure. hlorophyll ontent of Tsugru pples hrvested dys fter S- nd -fertilizer pplition. () hlorophyll ; () hlorophyll ; () totl hlorophyll; nd () hlorophyll / rtio. rs (mens ± SE, n = 4) with different letters differ signifintly (Tukey s HS test, p <.). 4 Journl of Food, griulture & Environment, Vol. (), pril 9

6 lorophyll (mg/g fresh wt) 4 hlorophyll (mg/g fresh wt). Totl lorophyll (mg/g fresh wt) Totl lorophyll (mg/g fresh wt) lorphyll / hlorophyll / Figure 9. hlorophyll ontent of pples hrvested dys fter S- nd fertilizer pplition. () hlorophyll ; () hlorophyll ; () totl hlorophyll; nd () hlorophyll / rtio. rs (mens ± SE, n = 4) with different letters differ signifintly (Tukey s HS test, t p <.). onlusions redued lipid peroxidtion nd inresed the totl ntioxidnt pity, phenolis, sori id, nthoynin nd hlorophyll ontents in the fruit peel of Tsugru nd pples, resulting in llevition of sunurn symptoms. These results suggest speifi -medited mehnisms tht ontriute to norml ell funtions under high temperture nd solr rdition stress. The results lso indite strong positive reltionship etween nd ntioxidnt levels, suggesting plys role in the tivtion of plnt defense mehnisms ginst sunurn. Hene, tretment ppers to e promising pproh to improve pple produtivity prtiulrly during the hot periods of the yer. Referenes wd, M.., Wgenmkers, P.S. nd e Jger,.. Effets of light on flvonoid nd hlorogeni id levels in the skin of Jongold pples. Si. Horti. :9-9. ergh, O., Frnken, J., Vn Zyl, E.J., Kloppers, F. nd empers,.9. Sunurn on pples: Preliminry results of n investigtion onduted during the 9/99 seson. eiduous Fruit Grower :-. hung, Y.., hen, S.J., Hsu,.K., hng,.t. nd hou, S.T.. Studies on the ntioxidtive tivity of Grptopetlum pr-guyense E. Wlther. Food hem. 9: ohen, S.., Trr, J.M. nd Kennedy, J... ssessing the impt of temperture on grpe phenoli metolism. nlyti hem. t 6:-6. Felietti,.. nd Shrder, L. E.. hnges in pigment onentrtions ssoited with the degree of sunurn rowning of Fuji pple. J. m. So. Hort. Si. :-4. 6 Go, Q. nd Zhng, L.. Ultrviolet--indued oxidtive stress nd ntioxidnt defense system responses in sorte-defiient vt mutnts of ridopsis thlin. J. Plnt Physiol. 6:-4. Gind, J. nd Wnd, S.J.E.. omprtive effets of evportive ooling, kolin prtile film nd shde net on the ontrol of sunurn nd fruit qulity in pples. Hort. Si. 4:9-96. Glenn,.M., Prdo, E., Erez,., MFerson, J. nd Puterk, G.J.. refletive proessed-kolin prtile film ffets fruit temperture, rdition refletion nd sunurn in pple. J. m. So. Hort. Si. : Hlliwell,. nd Gutteridge, J.M Lipid peroxidtion, oxygen rdils, ell dmge nd ntioxidnt therpy. Lnet :96 9. Hodges,.M., elong, J.M., Forney,.F. nd Prnge, R.K Improving the thiorituri id-retive-sustnes ssy for estimting lipid peroxidtion in plnt tissues ontining nthoynin nd other interfering ompounds. Plnt :64 6. Imsu, K., Sekozw, Y., Kmuro, Y., Sugy, S. nd Gemm, H.. Improvement of fruit qulity y S- nd the fertilizer formulted K, P, Mg,, Mn ontining S- s pre-hrvest pplition on pehes nd pples. t Horti..4:9-4. Kmuro, Y.. MIYOI; new fertilizer ontining sisi id. om. Pro. Intl. Plnt Prop. So. :6-. Lurie, S. nd Klein, J quisition of low temperture tolerne in tomtoes y exposure to high temperture stress. J. m. So. Hort. Si. 6:-. 4 M, F. nd heng, L.. The sun-exposed peel of pple fruit hs higher xnthophyll yle-dependent therml dissiption nd ntioxidnts of the sorte-glutthione pthwy thn the shded peel. Plnt Si. 6:9-. Merzlyk, M.N., Solovhenko,.E. nd hivkunov, O... Ptterns of pigment hnges in pple fruits during dpttion to high sunlight nd sunsld development. Plnt Physiol. iohem. 4: Muri, M.., Ver,. nd Gri-rmon, F. 99. Effet of proessing methods on spinh: Proximte omposition in ftty ids Journl of Food, griulture & Environment, Vol. (), pril 9 4

7 nd solule protein. J. Si. Food gri. 9:4-46. Nel,. nd lton, R. 9. Sunurn on Grnny Smith. eiduous Fruit Grower :64. Oster, U., Tnk, R., Tnk,. nd Rudiger, W.. loning nd funtionl expression of the gene enoding the key enzyme for hlorophyll iosynthesis (O) from ridopsis thlin. Plnt J. :-. 9 Pdmvti, M., Skthivel, N., Thr, K. V. nd Reddy,. R. 99. ifferentil sensitivity of rie pthogens to growth inhiition y flvonoids. Phytohemistry 46:499-. Pntelidis, G.E., Vsilikkis, M., Mngnris, G.. nd imntidis, G.. ntioxidnt pity, phenol, nthoynin nd sori id ontents in rsperries, lkerries, red urrnts, gooseerries nd ornelin herries. Food hem. :-. Prhomhuk, P. nd Meheriuk, M.996. Orhrd ooling with pulsed over tree irrigtion to prevent sunurn nd improve fruit qulity of Jongold pples. Hort. Si. :-4. Sirm, R.K., Srivstv, G.., nd Sxen,... Inresed ntioxidnt tivity under elevted temperture: mehnism of het stress tolerne in whet genotypes. iol. Plnt. 4:4-. Slisury, F.. nd Ross, Plnt Physiology. Wdsworth Pulishing ompny, In., elmont, liforni, US, 4 p. 4 Shrder, L.E., Zhng, J. nd uplg, W.K.. Two types of sunurn in pple used y high fruit surfe (peel) temperture. Plnt Helth Progress 4 (Otoer). Shrder, L.E., Zhng, J. nd Sun, J.. Environmentl stresses tht use sunurn of pple. t Horti. 6: Shupp, J.R., Fllhi, E. nd hun, I.J.. Effet of prtile film on fruit sunurn, mturity nd qulity of Fuji nd Honeyrisp pples. HortTehnol. :-9. Shinomiy, K., Tokung, S., Shigemoto, Y. nd Kmei,.. Effets of seed ot extrt of lk soyens on rdil mze performne in rts. lin. Exp. Phr. Physiol. : 6. Slinkrd, K. nd Singleton, V. L. 9. Totl phenol nlysis: utomtion nd omprison with mnul methods. m. J. Enol. Vit. : Viente,.R., ivello, P.M., Mrtinez, G.. nd hves,.r.. Qulity of het-treted strwerry fruit during refrigerted storge. Posthrvest iol. Tehnol. :9-. Wnd, S.J.E., Theron, K.I., kermn, J. nd Mris, S.J.S. 6. Hrvest nd post-hrvest pple fruit qulity following pplitions of kolin prtile film in South frin orhrds. Si. Horti. : 6. Wng, Y.S., Tin, S.P. nd Yong, X.. Effets of high oxygen onentrtion on pro-nd nti-oxidnt enzymes in peh fruits during posthrvest periods. Food hem. 9:99-4. Wunshe, J.N., Lomrdini, L. nd Greer,.H. 4. Surround prtile film pplitions effets on whole nopy physiology of pple. t Horti. 66:6-. Yhi, E.M., Soto-Zmor, G. J., reht, K. nd Grde,.. Posthrvest hot ir tretment effets on the ntioxidnt system in stored mture-green tomtoes. Posthrvest iol. Tehnol. 44:-. 4 Zheng, W. nd Wng, S.Y.. Oxygen rdil soring pity of phenolis in lueerries, rnerries, hokeerries, nd lingonerries. J. gri. Food hem. :-9. Zhung, H., Hildernd,.F. nd rth, M.M. 99. Temperture influened lipid peroxidtion nd deteriortion in rooli uds during posthrvest storge. Posthrvest iol. Tehnol. :49. 4 Journl of Food, griulture & Environment, Vol. (), pril 9