Bagging Treatment Influences Production of C 6 Aldehydes and Biosynthesis-Related Gene Expression in Peach Fruit Skin

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1 Molecules 2014, 19, ; doi: /molecules OPEN ACCESS molecules ISSN Article Bgging Tretment Influences Production of C 6 Aldehydes nd Biosynthesis-Relted Gene Expression in Pech Fruit Skin Ji-Yun Shen 1,2, Lei Wu 1, Hong-Ru Liu 1, Bo Zhng 1, *, Xue-Ren Yin 1, Yi-Qing Ge 3 nd Kun-Song Chen Lortory of Fruit Qulity Biology, The Stte Agriculture Ministry Lortory of Horticulturl Plnt Growth, Development nd Qulity Improvement, Zhejing University, Zijingng Cmpus, Hngzhou , Chin; E-Mils: sjy_sd@zju.edu.cn (J.-Y.S.); wulei.j@163.com (L.W.); her2008drem@163.com (H.-R.L.); xuerenyin@zju.edu.cn (X.-R.Y.); kun@zju.edu.cn (K.-S.C.) Physiologicl Lortory for South Chin Fruits, College of Horticulture, South Chin Agriculturl University, Gungzhou , Chin Chin Rurl Technology Development Center, Beijing , Chin; E-Mil: @163.com * Author to whom correspondence should e ddressed; E-Mil: ozhng@zju.edu.cn; Tel.: ; Fx: Received: 27 June 2014; in revised form: 20 August 2014 / Accepted: 25 August 2014 / Pulished: 29 August 2014 Astrct: Bgging is useful method to improve fruit qulity y ltering its exposure to light, wheres its effect on fruit voltiles production is inconsistent, nd the genes responsile for the oserved chnges remin unknown. In the present study, single-lyer yellow pper gs were used to study the effects of gging tretment on the formtion of C 6 ldehydes in pech fruit (Prunus persic L. Btsch, cv. Yulu) over two succeeding sesons. Higher concentrtions of n-hexnl nd (E)-2-hexenl, which re chrcteristic rom voltiles of pech fruit, were induced y gging tretment. After gging tretment, pech fruit hd significntly higher LOX nd HPL enzyme ctivities, ccompnying incresed contents of C 6 ldehydes. The gene expression dt otined through rel-time PCR showed tht no consistent significnt differences in trnscript levels of LOX genes were oserved over the two sesons, ut significntly up-regulted expression ws found for PpHPL1 fter gging tretment In ddition, gging-treted fruit produced more (E)-2-hexenl nd hd higher expression levels of PpHPL1 during posthrvest ripening t room temperture. The regultory role of the LOX-HPL pthwy

2 Molecules 2014, on the iosynthesis of n-hexnl nd (E)-2-hexenl in response to gging tretment during pech fruit development is discussed in the text. Keywords: gging; C 6 ldehydes; HPL; LOX; pech fruit 1. Introduction Voltile compounds intercting with sugrs nd cids contriute predominntly to fruit flvor. In pech fruit, more thn 100 voltile compounds hve een identified [1], nd green-note sensory voltiles, such s n-hexnl nd (E)-2-hexenl, contriute to the formtion of the chrcteristic pech rom [2,3]. The iosynthesis of voltile compounds is influenced y pech fruit genetics [4], developmentl stge [5], nd environmentl fctors, such s temperture [6,7], oxygen [8,9], nd light [10,11]. As one of the most populr cultivtion pproches, fruit gging is extensively prcticed in Chin nd Jpn to improve the fruit qulity y ltering its exposure to light. For exmple, fruit gging hs een selected s useful method for the study of nthocynin synthesis in fruits such s pple [12] nd per [13]. However, there is little informtion on the effects of gging on fruit voltiles, nd some reports re even contrdictory. In the cse of pech, gging with ornge pper cused no effect on the rom compounds of whole pech fruit t hrvest, ut significntly incresed the (E)-2-hexenl contents in the pech skin [10]. Another report showed tht n-hexnl nd 2-hexenl, s well s C 6 -relted esters, were significntly lower in pech fruit treted with two-lyered pper gs (lck inner nd rown outer ppers) compred with non-gged fruit [11]. These different results indicte the differentil influence of gging on the contents of C 6 ldehydes, ut the mechnism through which gging ffects the formtion of pech fruit voltiles remins uncler. Voltile n-hexnl nd (E)-2-hexenl re directly derived from the clevge of 13-hydroperoxide ctlyzed y hydroperoxidelyse (HPL); the 13-hydroperoxide is produced y the metolism of C 18 polyunsturted ftty cids, ctlyzed y lipoxygenses (LOXs) [14]. LOX cn ct on C 18 ftty cids t either the 9ꞌ or 13ꞌ position to generte two types of oxylipins nd is thus clssified s 9- nd 13-LOX. Accordingly, the HPL enzymes re minly ctegorized s 9- nd 13-HPL depending on their sustrte specificity [15], nd third type, nmely 9/13-HPL, which hs dul 9- nd 13-hydroperoxide specificity, hs lso een descried [16]. The dvnces in understnding of the roles of LOX nd HPL in fruit voltile iosynthesis hve een previously reviewed [17]. In our previous studies, we cloned four LOX genes nd one HPL gene in pech nd nlyzed the expression ptterns of these genes during posthrvest ripening t mient temperture nd fter storge t low temperture plus susequent shelf-life [18,19]. The ove descried reports provide clues tht cn e further used in the nlysis of gene expression nd enzyme ctivity to etter understnd the regultory role of gging on pech fruit ldehyde iosynthesis. In the present work, melting pech fruits were treted with gs over two succeeding sesons. First, the effect of gging on the concentrtions of the green-note n-hexnl nd (E)-2-hexenl ws nlyzed. Second, the gene expression ptterns of LOX nd HPL nd enzyme ctivities were studied fter gging tretment. Third, the effects of the gging tretment on the weight, colortion, esters

3 Molecules 2014, nd lctones t hrvest were discussed. Bsed on these findings, possile mechnism through which gging ffects voltile synthesis in pech fruit ws discussed. 2. Results nd Discussion Bgging tretment is n effective pproch tht is used to ffect the colortion nd flvor of fruits, including the rom-relted voltiles. Of these voltile compounds, C 6 ldehydes, such s n-hexnl nd (E)-2-hexenl, contriute to the green sensory notes of fruit [3]. Despite the importnce of these voltiles, the role of gging on the regultion of rom-relted voltiles remins uncler. As the chrcteristic voltile ldehydes for pech flvor, the chnges in n-hexnl nd (E)-2-hexenl in pech fruit in response to gging tretment were nlyzed. As shown in Tle 1, higher levels of n-hexnl were oserved in the pech fruits treted with yellow pper gs thn in the non-gged fruits, lthough differences were not significnt. The content of (E)-2-hexenl, which is nother C 6 ldehyde, ws pproximtely 60% higher in the gging-treted fruits thn in the controls in the 2010 seson, nd n ccumultion of pproximtely 3.7-fold ws found in the pper g-treted fruits in the 2011 seson (Tle 1). When the fruits were held t room temperture for 3 dys, contents of (E)-2-hexenl declined from μg g 1 FW to μg g 1 FW in pech fruits treted with g, while control fruits emitted lower levels during posthrvest ripening process (Figure 1). Tle 1. Effects of gging tretment on the concentrtions of n-hexnl nd (E)-2-hexenl (µg g 1 FW) in pech fruit skin t hrvest Fruit Seson 2011 Fruit Seson Control Bgged Control Bgged n-hexnl ± ± ± ± 5.03 (E)-2-Hexenl ± ± ± ± 1.86 Note: Ech vlue represents the men ± stndrd error of three replictes. Vlues indicted y different letters re significnt different to controls (LSD, 0.05). Figure 1. Chnges in (E)-2-hexenl contents nd PpHPL1 expressions of pech fruit skin during posthrvest ripening (2011 fruit seson). Ech vlue represents the men ± stndrd error of three replictes. The significnt differences re indicted with different letters (LSD, 0.05). Reltitve intensity (E)-2-hexenl (μg.g -1 ) Control Bgged dy 0 dy 3

4 Molecules 2014, Our results showed tht the gging tretment led to remrkly higher ccumultions of n-hexnl nd (E)-2-hexenl in pech fruit skin tissue over two succeeding sesons. The induction of C 6 ldehydes in response to gging tretment ws lso reported y Ji et l. [10], who oserved significntly higher undnce of (E)-2-hexenl in pech fruit skin tissue fter gging with triple prchment pper gs (50% sunlight trnsmission). In grpe fruit, sunlight shding lso significntly incresed the formtion of C 6 compounds [20]. However, in contrst to the ove results, pech fruit gging with two-lyered pper g (lck inner nd rown outer ppers) led to reduction in C 6 ldehydes [11], ut no informtion on the sunlight trnsmission of these pper gs ws provided. These inconsistencies could e due to the different sunlight trnsmission properties of the gs used. In ddition, fruit cultivrs nd mturity stges re possily other importnt fctors tht ffect voltile compound formtion in response to gging tretment. The LOX-HPL pthwy is considered to e involved in the iosynthesis of C 6 ldehydes [14]. To clrify the mechnism through which gging tretment increses the contents of C 6 ldehydes in pech fruit, the gene expression ws nlyzed to identify possile genes ssocited with the oserved chnges in ldehydes. Trnscripts of PpLOX1 nd PpLOX3 were induced y gging tretment, however, no significnt differences were oserved over two succeeding sesons (Figure 2). PpLOX2 exhiited different expression ptterns fter gging, ut the levels of difference were oth elow the 0.05 level of significnce. In the cse of nother memer PpLOX4, yellow pper g tretment cused no significnt inhiitions in expression levels either in 2010 or 2011 fruit seson (Figure 2). Although out four-fold higher LOX enzyme ctivities were induced fter gging (Figure 3), no significnt induction of trnscript levels were oserved for four LOX genes over two sesons, indicting the existence of other gene fmily memers tht contriute to chnges in the enzyme ctivity in response to gging tretment. Figure 2. Effect of gging tretment on the expression of PpLOXs nd PpHPL1 in pech fruit skin. Ech vlue represents the men ± stndrd error of three replictes. The significnt differences re indicted with different letters (LSD, 0.05). Reltive Intensity Reltive intensity seson 2011 seson Control Bgged PpLOX1 PpLOX2 PpLOX3 PpLOX4 PpHPL1 Genes

5 Molecules 2014, Figure 3. Effect of gging tretment on the concentrtions of C 6 ldehydes (A), LOX ctivity (B), nd HPL ctivity (C) in pech fruit skin (2010 fruit seson). Ech vlue represents the men ± stndrd error of three replictes. The significnt differences re indicted with different letters (LSD, 0.05). C6 ldehydes (μg g -1 FW) Enzyme ctivity (U mg -1 protein) A B C Control Bgged With respect to the HPL gene, the gging tretment significntly induced the ccumultion of PpHPL1 trnscripts of pech fruit in the 2010 seson, nd significnt ccumultion of expression levels were found in fruit treted with gs in the 2011 seson (Figure 2). Our results lso showed tht the induction of PpHPL1 expression ws ccompnied with n increse in the HPL enzyme ctivities. Compred with the non-gged control tch, the gging tretment induced n pproximtely 1.8-fold increse in the HPL enzyme ctivity (Figure 3). In greement with our previous study [18], expression levels of PpHPL1 tended to decrese during posthrvest ripening t room temperture (Figure 1). High expression levels of PpHPL1 in gged pech fruit were pproximtely times higher thn tht in the controls (Figure 1). In order to find more evidence to confirm the ssocition of HPL nd C 6 ldehydes, concentrtions of the voltile compounds nd expression ptterns of the gene were nlyzed during pech fruit mturtion. Significnt reduction of (E)-2-hexenl contents ws oserved during the fruit mturtion, decresing from μg g 1 t 106 DAB (endocrp hrden stge) to μg g 1 t 136 DAB (hrvest time) (Figure A1). QPCR results showed tht trnscripts of PpHPL1 tended to decline s fruit mturing (Figure A1). As expected, expression levels of PpHPL1 were generlly consistent with contents of n-hexnl nd (E)-2-hexenl during the pech fruit mturtion. Regression nlysis showed tht positive correltion R 2 = 0.83 ws oserved etween ldehydes contents nd HPL expression during pech mturtion. Our previous study showed tht PpHPL1 elong to 13-HPL group, nd ws suggested to puttively contriute to the formtion of voltiles during posthrvest ripening nd senescence [18]. Tken together, genes expression dtset indicted tht PpHPL1 ws likely ssocited with the chnges in the contents of C 6 ldehydes in fruit during pech development. Formtion of voltile compounds is influenced y fruit mturity nd ripening stges. Generlly, green-note voltiles tended to decrese while contents of fruity-note voltiles ccumulted during fruit ripening [17,18,21]. In our present study, the contents of hexyl cette, (Z)-3-hexenyl cette, nd (E)-2-hexenyl cette were slightly lower in the pech fruit treted with gs thn tht in the controls

6 Molecules 2014, (Tle 2). The gging-treted pech fruit lso produced reltively lower levels of γ-octlctone, γ-declctone nd δ-declctone (Tle 2). Although the levels of esters nd lctones were reduced in the gged fruit, they were generlly elow the 0.05 level of significnce. Tle 2. Effects of gging tretment on the concentrtions of esters nd lctones (µg g 1 FW) in pech fruit skin t hrvest. These effects were determined in two succeeding sesons. Voltiles 2010 Fruit Seson 2011 Fruit Seson Control Bgged Control Bgged Hexyl cette 2.28 ± ± ± ± 0.02 (Z)-3-Hexenyl cette 2.03 ± ± ± ± 0.03 (E)-2-Hexenyl cette 1.47 ± ± ± ± 0.09 γ-octlctone 3.13 ± ± 0.36 UD UD δ-declctone UD UD 0.37 ± ± 0.04 γ-declctone 0.41 ± ± ± ± 0.11 Note: Ech vlue represents the men ± stndrd error of three replictes. Vlues indicted y different letters re significntly different to controls (LSD, 0.05). UD, elow the determintion limit. In ddition to voltile compounds, differences in pech fruit mturity relted qulity were lso nlyzed. The weights of the pech fruits were not significntly ffected y gging tretment in oth fruit sesons (Tle 3). However, the fruit peel color ws significntly chnged fter gging tretment. The lightness (L) vlue of the pech fruit treted with yellow gs ws significntly higher thn tht otined for the control fruits (Tle 3). The non-gged pech fruits hd low hue ngle (H) nd chrom (C *) vlues, nd exhiited low contents of chlorophyll, chlorophyll, nd totl chlorophyll (Tle 3). Vlues of fruit firmness were slightly high in g treted pech fruit, while content of TSS nd flesh juiciness were slightly low in fruit treted with yellow g (Tle 3). Although these differences were generlly elow the 0.05 level of significnce, trends were pprent. The ove dtset showed tht pech fruit fter gging produced higher vlues of firmness, nd hd lower juiciness nd TSS content, indicting tht gging tretment influence the fruit mturity. Tle 3. Effects of gging tretment on the weight (g), colortion (L, H, C), chlorophyll (µg g 1 FW), firmness, TSS ( Brix) nd juiciness (%) of pech fruits t hrvest Fruit Seson 2011 Fruit Seson Control Bgged Control Bgged Weight ± ± ± ± 4.98 L ± ± ± ± 1.22 H ± ± ± ± 0.68 C * ± ± ± ± 0.33 Chlorophyll 2.95 ± ± 0.35 NA NA Chlorophyll 1.77 ± ± 0.41 NA NA Firmness ± ± 2.69 NA NA TSS ± ± 0.45 NA NA Juiciness ± ± 2.44 NA NA Note: Ech vlue represents the men ± stndrd error of three replictes. Vlues indicted y different letters re significntly different to controls (LSD, 0.05). NA mens dt not ville.

7 Molecules 2014, Recently, more nd more reports hve shown tht tretment with different light could effectively influence the emissions of plnt voltiles [22 24], ut the moleculr sis for this remins uncler. Relese of pech drft genome sequence [25] is going to provide n opportunity to clone nd identify more genes, thus to understnd mechnism of the effects of light on voltiles formtion. In ddition, it will e interesting to determine whether chnges in voltile compounds ffected y gging tretment could ffect humn perception of flvor qulity. 3. Experimentl 3.1. Plnt Mterils nd Smpling Melting pech fruits (Prunus persic L. Btsch cv. Yulu) were used in this study. The experiments were conducted over two succeeding sesons (2010 nd 2011) in the sme commercil orchrd in Ningo, Zhejing Province, Chin, where gging fruit with single-lyer yellow g is used s cultivtion method to protect Yulu pech fruit. Twenty pech trees were divided into two groups sed on the gging tretment in the 2010 fruit seson. Ten fruits of ech tree were rndomly selected nd covered with single-lyer yellow pper gs (15.7 cm 14.7 cm, Zhungnongyuguo Compny, Ningo, Chin) on July 19th. The non-gged pech fruits were exposed to direct sunlight nd used s control. In the 2011 fruit seson, nine pech trees of ech of ten fruit were covered with single-lyer yellow pper gs during the second stge of fruit development (period of slow growth) on 25 July, nd fruits from nine other pech plnts were not gged nd used s controls. The sunlight trnsmission nlysis using luminometer (TES-1339, TES Electricl Electronic Corp., Tipei, Tiwn) showed tht the sunlight trnsmission through single-lyer yellow pper g is out 25%. Both the gged nd the non-gged pech fruits were hrvested t commercil mturity (firm-mture stge) over two sesons. To void their exposure to light efore the qulity nlysis, the gged fruits were hrvested without removing the gs. The fruits were trnsported to the lortory on the dy of hrvest nd screened for uniform size nd freedom from defects nd wounds. In the second fruit seson, pech fruit smpled on 11 July, 25 July, 1 August, 5 August nd 10 August, which represent 106, 120, 127, 131 nd 136 dys fter loom (DAB) respectively. Moreover, fruit hrvested t 136 DAB were lso llowed for posthrvest rtificil ripening t room temperture for 3 dys. For iochemicl nd moleculr nlyses, three replictes of five fruit ech were used in ech seson Fruit Qulity Evlution The weight nd peel color of the fruits were mesured on the dy of hrvest nd re shown in Tle 1. The fruit peel color, which is expressed through the lightness (L), hue ngle (H ), nd chrom (C *), ws mesured on oth sides of the equtor of the pech fruit. The lightness ws mesured directly, nd the hue ngle nd chrom were clculted ccording to the methods descried in previous study [26]. Chlorophyll ws extrcted from fruit peel s previously descried y Lewndowsk et l. [27] with some modifictions. Fruit peel (200 mg) ws ground nd homogenized in 90% cold cetone (4 ml) nd centrifuged t 10,000 g for 20 min t 4 C. The sornce of the superntnt ws mesured using DU800 spectrophotometer (Beckmn, Fullerton, CA, USA) t 644 nd 662 nm. The content of

8 Molecules 2014, chlorophyll (Chl ) nd chlorophyll (Chl ) were clculted using following equtions: [Chl ] = 9.78A A 644 nd [Chl ] = 21.40A A 662. A TA-XT2i plus texture nlyzer (Stle Micro System, Godlming, Englnd) equipped with 7.9 mm dimeter hed ws pplied for fruit firmness mesurement s descried y Zhng et l. [18]. Two mesurements were mde on opposite sides of ech fruit fter the removl of 1 mm thick skin slice, nd dt were expressed s newtons (N). Totl solule solids (TSS) were mesured y slicing oth ends of ech fruit [18]. Three drops of juice from ech slice were then pplied to n PR-101α digitl hnd-held refrctometer (Atgo, Tokyo, Jpn). TSS dt were expressed in degrees Brix. For juice mesurements, 4 flesh cues were tken from ech fruit nd the weight of juice relesed y the pressure produced from texture nlyzer (TA-XT2i Plus) with 10 cm dimeter hed ws determined. The crushed flesh remining ws lso weighed, nd the proportion of juice in reltion to the totl flesh (mss/mss) ws clculted nd expressed s %, method previously descried y Zhng et l. [19]. The fruit peel tissue (pproximtely 1-mm-thick) ws sliced using peeler, frozen in liquid nitrogen, nd stored t 80 C until further use. Three replictes of ech of five fruits were used for the voltile nd moleculr nlyses Fruit Voltile Anlysis The voltile compounds in pech fruit were mesured using the method descried y Zhng et l. [18]. Frozen pech peel tissue ws ground into fine powder in liquid nitrogen nd one grm ws trnsferred to 15 ml glss vil contining sturted NCl solution. Then, 2-octnol (30 µl, Fluk, St. Louis, MO, USA) ws dded to the lend s n internl stndrd, nd mixture ws vortexed for 10 s. The smples were equilirted for 30 min nd sujected to mnul solid-phse micro-extrction nlysis (SPME) with fier coted with 65 µm of polydimethylsiloxne nd divinylenzene (Supelco Co., Bellefonte, PA, USA). Susequently, the voltile compounds were nlyzed on n Agilent 6890N GC equipped with DB-WAX column (0.32 mm, 30 m, 0.25 µm, J&W Scientific, Folsom, CA, USA). The chromtogrphy temperture ws progrmmed s follows: the oven ws initilly mintined t 34 C for 2 min, incresed to 60 C t rte of 2 C min 1, incresed to 220 C t rte of 5 C min 1, nd mintined t 220 C for 2 min. Nitrogen ws used s crrier gs t flow rte of 1.0 ml min 1. Using the internl stndrd vlue s the reference vlue, the quntittive vlues of the voltile compounds were clculted sed on the stndrd curves of the uthentic compounds supplied y Sigm-Aldrich (St. Louis, MO, USA) Enzymes Activity Anlysis Anlysis of LOX ctivity ws performed y the method shown y Echeverri et l. [28]. Powdered peel tissue (100 mg) tken from three peches ws homogenized in extrction solution (1 ml). The extrction solution ws consisted of 2 mm dithiothreitol, 1 mm EDTA, 0.1% (v/v) Triton X-100, 1% (w/v) PVPP nd 0.1 M phosphte (ph 7.5). The sustrte solution ws consisted of 8.6 mm linoleic cid, 0.25% (v/v) Tween-20, 10 mm NOH nd 0.1 M phosphte (ph 8.0). LOX ctivity in the crude extrct ws mesured y following spectrophotometriclly the increse in sornce t 234 nm. One

9 Molecules 2014, ctivity unit (U) ws defined s the increse in one unit of sornce t 234 nm per minute, nd results were expressed s specific ctivity (U mg protein 1 ). The crude HPL enzyme ws extrcted ccording to the method descried y Fukushige nd Hildernd [29]. The extrction uffer consisted of 100 mm Tris HCl (ph 8.5), 3 mm EDTA, 3 mm dithiothreitol, 0.5% (w/v) Triton X-100, 1% (w/v) protese inhiitor cocktil, nd 5 g L 1 PVPP. One grm of peel tissue ws ground nd dded into 2 ml of extrction uffer, nd the homogente ws centrifuged t 12,000 rpm t 4 C. The enzyme rection occurred in the presence of NADH nd yest lcohol dehydrogense, where the ldehyde ws converted to lcohol y consuming NADH. The reduction of NADH ws spectrophotometriclly mesured y the loss sornce t 340 nm. The protein content ws mesured [30], nd the HPL enzyme ctivity ws expressed s U mg protein RNA Extrction nd Gene Expression Anlysis The totl RNA ws extrcted from one grm of frozen peel tissue ccording to the method descried y Zhng et l. [31]. RNse-free DNse I (Ferments, St. Leon-Rot, Germny) ws used to remove ny possile genomic DNA. The first-strnd cdna synthesized from 3 µg of DNA-free RNA ws susequently used for the gene expression nlysis of PpLOXs nd PpHPL1 using CFX96 rel-time PCR instrument (Bio-Rd, Hercules, CA, USA). The primer sequences of ll of the genes ssyed nd the temperture progrm for the rel-time PCR ssy re shown in our previous study [18]. At lest three different RNA isoltions nd cdna syntheses were used s replictes for the gene expression nlysis. The expression levels were expressed s rtio reltive to the levels found in non-gged fruit, which were set to 1. For pech smpled t differentil ripening stges, the first time point (106 DAB) expressions were set to Sttisticl Anlysis The experiment ws designed through completely rndomized design. Dt from ech tretment nd control were sujected to nlysis of vrince (significnce t 0.05) nd mens were seprted using lest significnt difference (LSD). The significnt differences re indicted in the figures nd tles with different letters. The sttisticl nlysis nd figures were generted using OriginPro 9.0 (OriginL Corportion., Northmpton, MA, USA). 4. Conclusions In summry, gging tretment of pech fruit with single-lyer yellow pper gs produced more contents of n-hexnl nd (E)-2-hexenl in skin tissues vi inducing PpHPL1 expression nd enzyme ctivity. Our results indicte tht covering the fruits with different gs with vrious sunlight trnsmission properties is potentil effective method for the ltertion of fruit voltile compounds. Acknowledgments This work ws supported y the Ntionl Science Foundtion of Chin ( ), the Ntionl High Technology Reserch nd Development Progrm of Chin (2012AA101702), the Ntionl Key

10 Molecules 2014, Technology Reserch nd Development Progrm of the Ministry of Science nd Technology of Chin (2013BAD19B05). Author Contriutions Ji-yun Shen crried out gene expression nd mnuscript drft preprtion, Lei Wu performed GC nd dt nlysis, Hong-ru Liu crried out gging tretment nd fruit qulity nlysis, Bo Zhng designed experiment nd prepred the mnuscript, Xue-ren Yin crried out dt nlysis, Yi-qing Ge gve criticl revision of the mnuscript, nd Kun-song Chen prepred the mnuscript. Appendix Figure A1. Chnges in n-hexnl, (E)-2-hexenl contents, nd PpHPL1 expression levels of pech fruit skin during mturtion. Men vlue of three replictes is shown with stndrd error, nd different letters ove the symols indicte significnt differences (LSD, 0.05). Contents (μg g -1 FW) Reltive intensity n-hexnl (E)-2-hexenl c PpHPL1 c c Dys fter loom Conflicts of Interest The uthors declre no conflict of interest. References 1. Auert, C.; Milhet, C. Distriution of the voltile compounds in the different prts of white-fleshed pech (Prunus persic L. Btsch). Food Chem. 2007, 102, Horvt, R.J.; Chpmn, G.W.; Roertson, J.A.; Meredith, F.I.; Scorz, R.; Cllhn, A.M.; Morgens, P. Comprison of the voltile compounds from severl commercil pech cultivrs. J. Agric. Food Chem. 1990, 38, Edurdo, I.; Chieter, G.; Bssi, D.; Rossini, L.; Vecchietti, A. Identifiction of key odor voltile compounds in the essentil oil of nine pech ccessions. J. Sci. Food Agric. 2010, 90,

11 Molecules 2014, Montero-Prdo, P.; Bentye, K.; Nerin, C. Pttern recognition of pech cultivrs (Prunus persic L.) from their voltile components. Food Chem. 2013, 138, Sánchez, G.; Venegs-Cleron, M.; Sls, J.J.; Monforte, A.; Bdenes, M.L.; Grnell, A. An integrtive omics pproch identifies new cndidte genes to impct rom voltiles in pech fruit. BMC Genomics 2013, 14, Xi, W.P.; Zhng, B.; Shen, J.Y.; Sun, C.D.; Xu, C.J.; Chen, K.S. Intermittent wrming llevited the loss of pech fruit rom-relted esters y regultion of AAT during cold storge. Posthrvest Biol. Technol. 2012, 74, Cno-Slzr, J.; López, M.L.; Crisosto, C.H.; Echeverri, G. Voltile compound emissions nd sensory ttriutes of Big Top nectrine nd Erly Rich pech fruit in response to pre-storge tretment efore cold storge nd susequent shelf-life. Posthrvest Biol. Technol. 2013, 76, Ortiz, A.; Grell, J.; López, M.L.; Echeverri, G.; Lr, I. Voltile ester-synthesising cpcity in Trdielle pech fruit in response to controlled tmosphere nd 1-MCP tretment. Food Chem. 2010, 123, Yng D.S.; Blndrn-Quintn, R.R.; Ruiz, C.F.; Toledo, R.T.; Kys, S.J. Effect of hyperric, controlled tmosphere, nd UV tretments on pech voltiles. Posthrvest Biol. Technol. 2009, 52, Ji, H.J.; Arki, A.; Okmoto, G. Influence of fruit gging on rom voltiles nd skin colortion of Hkuho pech (Prunus persic Btsch). Posthrvest Biol. Technol. 2005, 35, Wng, Y.J.; Yng, C.X.; Liu, C.Y.; Xu, M.; Li, S.H.; Yng, L.; Wng, Y.N. Effects of gging on voltiles nd polyphenols in Wnmi peches during endocrp hrdening nd finl fruit rpid growth stges. J. Food Sci. 2010, 75, S455 S Liu, Y.L.; Che, F.; Wng, L.X.; Meng, R.; Zhng, X.J.; Zho, Z.Y. Fruit colortion nd nthocynin iosynthesis fter g removl in non-red nd red pples (Mlus domestic Borkh.). Molecules 2013, 18, Hung, C.H.; Yu, B.; Teng, Y.W.; Su, J.; Shu, Q.; Cheng, Z.Q.; Zeng, L.Q. Effects of fruit gging on coloring nd relted physiology, nd qulities of red Chinese snd pers during fruit mturtion. Sci. Hortic. 2009, 121, Schw, W.; Dvidovich-Riknti, R.; Lewinsohn, E. Biosynthesis of plnt-derived flvor compounds. Plnt J. 2008, 54, Mtsui, K. Green lef voltiles: Hydroperoxidelyse pthwy of oxylipin metolism. Curr. Opin. Plnt Biol. 2006, 9, De Domenico, S.; Tsesmetzis, N.; di Snsestino, G.P.; Hughes, R.K.; Csey, R.; Sntino, A. Sucellulr loclistion of Medicgo trunctul 9/13-hydroperoxide lyse revels new loclistion pttern nd ctivtion mechnism for CYP74C enzymes. BMC Plnt Biol. 2007, 7, Defilippi, B.G.; Mnriquez, D.; Luengwili, K.; Gonzlez-Aguero, M. Arom voltiles: Biosynthesis nd mechnisms of modultion during fruit ripening. Adv. Bot. Res. 2009, 50, Zhng, B.; Shen, J.Y.; Wei, W.W.; Xi, W.P.; Xu, C.J.; Ferguson, I.; Chen, K.S. Expression of genes ssocited with rom formtion derived from the ftty cid pthwy during pech fruit ripening. J. Agric. Food Chem. 2010, 58,

12 Molecules 2014, Zhng, B.; Xi, W.P.; Wei, W.W.; Shen, J.Y.; Ferguson, I.; Chen, K.S. Chnges in rom-relted voltiles nd gene expression during low temperture storge nd susequent shelf-life of pech fruit. Posthrvest Biol. Technol. 2011, 60, Bureu, S.M.; Rzungles, A.J.; Bumes, R.L. The rom of Musct of Frontignn grpes: Effect of the light environment of vine or unch on voltiles nd glycoconjugtes. J. Sci. Food Agric. 2000, 80, Kkiuchi, N.; Omiy, A. Chnges in the composition nd content of voltile constituents in pech fruits in reltion to mturity t hrvest nd rtificil ripening. J. Jpn. Soc. Hortic. 1991, 60, Bertoli, A.; Lucchesini, M.; Mensuli-Sodi, A.; Leonrdi, M.; Doveri, S.; Mgnosco, A.; Pistelli, L. Arom chrcteriztion nd UV elicittion of purple sil from different plnt tissue cultures. Food Chem. 2013, 141, Flr, V.; Amrsinghe, R.; Poldy, J.; Pichersky, E.; Brrow, R.A.; Pekll, R. The production of key florl voltile is dependent on UV light in sexully deceptive orchid. Ann. Bot. 2013, 111, Colquhoun, T.A.; Schwietermn, M.L.; Gilert, J.L.; Jwoski, E.A.; Lnger, K.M.; Jones, C.R.; Rushing, G.V.; Hunter, T.M.; Olmsted, J.; Clrk, D.G.; et l. Light modultion of voltile orgnic compounds from petuni flowers nd selected fruits. Posthrvest Biol. Technol. 2013, 86, Verde, I.; Aott, A.G.; Sclrin, S.; Jung, S.; Shu, S.; Mrroni, F.; Zheentyyev, T.; Dettori, M.T.; Grimwood, J.; Cttonro, F.; et l. The high-qulity drft genome of pech (Prunus persic) identifies unique ptterns of genetic diversity, domestiction nd genome evolution. Nt. Genet. 2013, 45, Infnte, R.; Frcuh, M.; Meneses, C. Monitoring the sensoril qulity nd rom through n electronic nose in peches during cold storge. J. Sci. Food Agric. 2008, 88, Lewndowsk, M.; Jrvis, P.G. Chnges in chlorophyll nd crotenoid content, specific lef re nd dry weight frction in sitk spruce, in response to shding nd seson. New Phytol. 1977, 79, Echeverri, G.; Grell, J.; López, M.L.; Lr, I. Voltile production, qulity nd rom-relted enzyme ctivities during mturtion of Fuji pples. Posthrvest Biol. Technol. 2004, 31, Fukushige, H.; Hildernd, D.F. Wtermelon (Citrullus lntus) hydroperoxide lyse gretly increses C-6 ldehyde formtion in trnsgenic leves. J. Agric. Food Chem. 2005, 53, Brdford, M.M. Rpid nd sensitive method for quntittion of microgrm quntities of protein utilizing principle of protein-dye inding. Anl. Biochem. 1976, 72, Zhng, B.; Chen, K.S.; Bowen, J.; Alln, A.; Espley, R.; Kruniretnm, S.; Ferguson, I. Differentil expression within the LOX gene fmily in ripening kiwifruit. J. Exp. Bot. 2006, 57, Smple Avilility: Smples of the pech fruit re ville from the uthors y the uthors; licensee MDPI, Bsel, Switzerlnd. This rticle is n open ccess rticle distriuted under the terms nd conditions of the Cretive Commons Attriution license (

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