grape pomace 葡萄

Invited review

Functional components of grape pomace:their composition,biological properties and potential applications

Jianmei Yu 1*&Mohamed Ahmedna 2

1Department of Family and Consumer Sciences,North Carolina A&T State University,1601East Market Street,Greensboro,NC 27411,USA

2Centre for Excellence in Post-Harvest Technologies,North Carolina A&T State University,500Laureate Way,Suite 4222,North Carolina Research Campus,Kannapolis,NC 28081,USA

(Received 20April 2012;Accepted in revised form 25July 2012)

Summary

The roles of functional foods on human health have been realised by more and more researchers,food producers and consumers.Functional food ingredients from both plant and animal sources such as die-tary ?bre,soy protein isolate,whey protein isolate and omega 3fatty acid have been widely used in func-tional food product development.Many fruit processing by-products such as grape,apple and orange peels are rich in bioactive phytochemicals,dietary ?bre and unsaturated fatty acids,hence have potential to serve as functional food ingredients.In this review,we summarise recent advancement of research in grape pomace (GP),the residual of grapes after wine making.The polyphenol pro?le of GP and their bio-logical,antioxidant and antimicrobial activities,the stability of GP polyphenols in food system,the inter-action between GP polyphenol and other food ingredients,as well as the functionalities of grape seed oil and GP ?bre are covered.

Keywords

Biological properties,dietary ?bre,grape pomace,grape seed oil,grape seed protein,polyphenol composition,thermal stability.

Introduction Grape pomace (GP)is a by-product of wine industry.GP consists mainly of peels (skins),seeds and stems and accounts for about 20–25%of the weight of the grape crushed for wine production.Grape seed is rich in extractable phenolic antioxidants such as phenolic acid,?avonoids,procyanidins and resveratrol,while grape skins contain abundant anthocyanins.The health bene?ts of GP polyphenols have been the great interest of researchers,food industry and nutraceutical industry.In addition to phenolic antioxidants,GPs also contain signi?cant amount of lipid,proteins,non-digestible ?bre and minerals.Grape seeds contain 13–19%oil,which is rich in essential fatty acids,about 11%protein,60–70%of non-digestible carbohydrates,and non-phenolic antioxidants such as tocopherols and beta-carotene (Rao,1994;Baydar &Akkurt,2001;Bravi et al.,2007;Llobera &Can ellas,2007).This review summarises the recent studies on major components of GP,their important properties and their possible applications that are.

Phenolic compounds of GP and their properties

Polyphenol composition of GP

Phenolics are the secondary metabolites of plants.Chemically,phenolics can be de?ned as substances pos-sessing an aromatic ring bearing one or more hydroxyl groups,including their functional derivatives (Shahidi &Naczk,2004).Polyphenols are compounds that have more than one phenolic hydroxyl group attached to one or more benzene rings (Vermerris &Nicholson,2006).Most food phenolics have more than one hydroxyl group attached on the aromatic ring;therefore,in this review,phenolics and polyphenols are used interchange-ably.In food science research,natural phenolics are generally classi?ed into classes and sub-classes based on the similarity of their chemical structures,that is,the types of building blocks that appear as repeated units.Four major classes of polyphenols found in foods are phenolic acids,?avonoids,lignans and stilbenes (Spencer et al.,2008).Major stilbenoids found in foods of plant origin are resveratrol and it glycosides.Resveratrol is a phytoalexin produced in the plant in response to pathogen attack.It has a low toxicity in humans and is a naturally occurring fungicide.

*Correspondent:Fax:3363347239;e-mail:jyu@https://www.wendangku.net/doc/ea12384708.html,

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Food Science and Technology 2013,48,221–237doi:10.1111/j.1365-2621.2012.03197.x

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Phenolic acids are phenols that possess one carbox-ylic acid functional group and are divided into hydroxycinnamic acids and hydroxybenzoic acids.The hydroxycinnamic acids are more common than hydroxy-benzoic acids,and they mainly include gallic acid,p -coumaric,ca?eic,chlorogenic acid,ferulic and sinapic acids.These acids are rarely found in the free form,except in food that has undergone freezing,sterilisation or fermentation.The bound forms are glycosylated derivatives or esters of quinic acid,shikimic acid and tartaric acid (Vermerris &Nicholson,2006).

The largest and best studied polyphenols are the ?avonoids.Based on their molecular structures,?avonoids are divided into seven subclasses:?avones,?avanones,?avonols,iso?avones,anthocyanidins/an-thocyanins,?avanols (or catechins and procyanidins)and chalcones (Karakaya,2004).Another group of ?avonoids,which are not included in this classi?ca-tion,are proanthocyanidins,also called,procyanidins,condensed tannins or oligomeric procyanidins (Prior &Gu,2005).

Although phenolics are present virtually in all plant foods,some fruits such as grape,apple,blueberry and cranberry are extremely rich in these bioactive com-pounds.In grape berries,the phenolic compounds reside mainly in the skins,seeds and short stems (Rodriguez et al.,2006;Poudel et al.,2008).GP is rich in extractable phenolic antioxidants (10–11%of dry weight)(Makris et al.,2007).Anthocyanins,catechins,procyanidins,?avonol glycosides,phenolic acids and stilbenes are the principal phenolic constituents found in GP (Montealegre et al.,2006).

Polyphenol composition of GP is variety dependant.The red varieties are usually rich in anthocyaninins that are neglected in while white varieties.Within the same variety,di?erent part has signi?cantly di?erent polyphenol composition.Cantos et al.(2002)analysed the polyphenol composition of four red and three white table grape varieties by HPLC-ADA-MS and found that anthocyanins were the main phenolics in red grapes ranging from 69(Crimson Seedless)to 151(Flame Seedless)mg kg à1fresh weight of grapes,whereas ?avan-3-ols were the most abundant phenolics in the white varieties ranging from 52(Dominga)to 81(Moscatel Italica)mg kg à1fresh weight of grapes.Flavan-3-ols were also detected and were identi?ed as gallocatechin,procyanidin B1,procyanidin B2,procy-anidin B4,procyanidin C1,catechin and epigallocate-chin.The study of the phenolic compounds content and antioxidant activity of pomace from the vini?cation of grape varieties widely produced in Brazil (Cabernet Sauvignon,Merlot,Bordeaux and Isabel)found that catechin was the most abundant non-anthocyanic com-pound identi?ed in the GP (150.16mg 100g à1)for all varieties;Cabernet Sauvignon pomace had the highest content of total phenolic compounds (75mg g à1),while Bordeaux variety showed the highest content of total anthocyanins (Rockenbach et al.,2011).

Phenolic compound distribution in di?erent parts of grapes was reviewed by Xia et al.(2010).The grape skins,part of pomace,are proven to be rich sources of anthocyanins,hydroxycinnamic acids,?avanols and ?avonol glycosides,whereas gallic acid and ?avanols were mainly present in the seed portion (Kammerer et al.,2004;Xia et al.,2010).Thirty-seven anthocya-nins have been separated by McCalluma et al.(2007)from Concord grape skin,and among them twenty-?ve were identi?ed using LC-MS.Among the anthocyanins identi?ed in GP,?ve of them are the 3-O -monogluco-sides of delphinidin,cyaniding,petunidin,peonidin and mavidin;another ?ve of them are the acetylgluco-sides of the ?ve anthocyandins.Malvidin 3-O -gluco-side was found to be the predominant anthocyanin.Grape seeds are rich in monomeric phenolic com-pounds,such as (+)-catechins,(à)-epicatechin and (à)-epicatechin-3-Ogallate,and dimeric,trimeric and tet-rameric procyanidins.Anthocyanin content of GP var-ies with wine vini?cation method and contact time.The longer the contact time,the lower the anthocyanin

content remains in the pomace (Go

mez-Plaza et al.,2006).

The phenolic pro?le of grape skin and seeds of European variety Vitis Vinifera has been intensively studied by scientists all over the world.Montealegre et al.(2006)quanti?ed many phenolic compounds in seeds and skins of ten V.vinifera grapes grown in the warm climate of Spain,including six white varieties (Chardonnay,Sauvignon blanc,Moscatel,Gewu rztr-aminer,Riesling and Viogner)and four red grape vari-eties (Cencibel,Cabernet Sauvignon,Merlot and Shiraz).They found that the skin of Viogner had the least of total hydroxycinnamates,catechin and procy-anidin dimmers,while that of Moscatel had the most of total hydroxycinnamates and ?avonols.Chardon-nay and Gewu rztraminer skins were the richest in cate-chin and procyanidins among white grape varieties.Red grape skin contains higher amount of hydroxycin-namates than white grape skin (31–55mg kg à1fresh grape),but much lower amount of catechins (9.0–18mg kg à1fresh grape),procyanidin dimmers (8.0–18mg kg à1fresh grape)and total ?avonols (12–19mg kg à1fresh grape).

The polyphenol composition of each part of the GP varies depending on the varieties of grapes and is in?u-enced by the growing location,climate,maturity and the time of fermentation (Fuleki &Ricardo da Silva,1997;Kennedy et al.,2000;Shi et al.,2003a;Monteal-egre et al.,2006).The per-berry extractable yield of all polyphenols decreased with maturity and followed sec-ond-order kinetics.The ?avan-3-ol monomers decreased most rapidly,followed by the procyanidin extension units and ?nally the terminal units.The

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relative proportion of procyanidin extension units did not vary with maturity(Kennedy et al.,2000).The phenolic composition of white and red grape seeds is comparable,but catechin,epicatechin and procyanidin B1are higher in white grape seeds.Overall,grape seeds contained lower amount of phenolic acid than grape skin,but rich in catechins and procyandins. Castillo-Mun oz et al.(2010)investigated the?avonol pro?les of white GPs from twenty-one V.vinifera white grape cultivars in Spain and found that?avonol pro?les of white grapes are dominated by quercetin-type?avonols,but some cultivars(e.g.Pedro Xime nez, Gewu rztraminer,Verdejo,Albillo,and Riesling)were characterised by relatively high and signi?cantly di?er-ent proportions of isorhamnetin-type?avonols.

The phenolic compounds in grape seeds are essen-tially all?avonoids,particularly,?avan-3-ols(catechin, epicatechin and epicatechin-3-O-gallate monomers) and their polypmers.Flavan-3-ols easily condenses into oligomeric procyanidins and polymeric com-pounds(condensed tannins).The dimeric procyanidins are often referred as B-series,and the trimeric procy-anidins as C-series.Five di?erent dimers(procyanidin B1,B2,B3,B4and B5)and two trimers(C1and C2) were identi?ed from grape skin and seeds(Shi et al., 2003a).Analysis of procyanidins extracted from both white and red grape seeds by electrospray ionisation–mass spectrometry(ESI-MS and ESI-MS/MS)in the positive mode found protonated molecules of procy-anidin species of nongalloylated and monogalloylated type-A and type-B oligomers,with degree of polymeri-sation2–5,and digalloylated oligomers,with degree of polymerisation2–3(Passos et al.,2007).

The native grape variety grown in the United States is Muscadine,which includes the popular cultivars of Carlos,Cowart,Jumbo,Magnolia,Sterling,Nesbitt, Scuppernong and Noble.These grapes have tougher skins and less seeds than V.vinifera grapes.The study conducted in our laboratory demonstrated that Mus-cadine GP had about5%more skin,but8%less seeds than Cabernet GP.The polyphenol composition of Muscadine GP and Cabernet GP was also di?erent. The contents of total extractable polyphenol,total anthocyanin and total?avonoid were36.42,0.88and 21.02mg gà1DM in Muscadine seeds,19.39,8.15 and4.78mg gà1DM in Muscadine skin,respectively (Yu et al.,2011).Ellagic acid,gallic acid,(à)-epicate-chin,(à)-epigallocatechin,catechin,myricetin,querce-tin,and kaempferol and some anthocyanidins including delphinidin,cyanidin,petunidin,peonidin and malvidin were identi?ed in the extract of Musca-dine pomace extract using the combination of reten-tion time and spectral properties on a reverse-phase HPLC–PDA(Wang et al.,2010).Approximately90% of the total anthocyanins in Muscadine grapes were 3,5-diglucoside of delphinidin,cyanidin and petunidin;the remaining10%were3,5-diglucoside of peonidin and malvidin;purple-skinned muscadine grapes(Jumbo and Cowart)have signi?cantly higher levels of anthocy-anins than bronze-skinned muscadine grapes(Carlos and Higgin)(Huang et al.,2009).

Resveratrol is another important polyphenol found in grape skins and seeds.Resveratrols are found lar-gely in the skins of red grapes and in other foods such as mulberries and peanuts(Sanders et al.,2000).The trans-resveratrol content was found to be 1.11–12.3 mg per100dry mass in grape skin,8.64±4.5mg per 100dry mass in white grape skin and1.42±0.18mg per100g dry mass in white grape seeds(Kammerer et al.,2004).The resveratrol content in grapes di?ers according to the variety of grape(Ector et al.,1996) and the grape maturation(Moreno et al.,2008).Mus-cadine grapes and their products were reported to con-tain more resveratrol than any other type of grape (Ector et al.,1996).‘Carlos’and‘Magnolia’Musca-dine cultivars had the greatest skin resveratrol concen-tration of all the Muscadine cultivars evaluated. Except for‘Sweet Jenny’,bronze cultivars had greater skin resveratrol concentration than black skinned culti-vars.‘Miss Blanc’Vitis labrusca grape had greater skin resveratrol concentration than all other cultivars (LeBlanc,2006).Resveratrol contents in grape tissues can be modi?ed by post-harvest technologies.Cold storage alone doubled skin stilbene concentration in ‘Carlos’grape,but UV irradiation did not signi?cantly change stilbene levels.In contrast,UV irradiation increased skin stilbene concentration by50%in ‘Noble’grape,but cold storage alone had no e?ect (LeBlanc,2006).Although certain amount of resvera-trols in grape transfers into wine during grape macera-tion,signi?cant amount of resveratrols remains in the pomace(Feijo o et al.,2008).

Like other plant materials,GP contains relatively higher amount of non-extractable polyphenols(NEP). Although the total polyphenol content in dry GP is about 4.8–5.4%(Makris et al.,2007),only2%of polyphenols in GP is extractable under mild conditions commonly used to develop polyphenol databases (Bravo&Saura-Calixto,1998).The majority portion of GP polyphenols has been reported to be highly polymerised condensed tannin,and some polyphenols form complex with?bre and are non-extractable unless strong acidic treatments are applied(Arranz et al., 2010).Monomeric and oligomeric proanthocyanidins are certainly soluble in the organic solvents usually used for polyphenol extraction,but a major propor-tion of high-molecular-weight proanthocyanidins and polyphenols complexed with protein or cell wall poly-saccharides remain insoluble(Huemmer&Schereier, 2008).The quanti?cation of NEP in GP needs hydro-lysis of GP residual to release the bound phenolics from cell wall or protein after soluble polyphenols are

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extracted (Ignat et al.,2011).The NEP content of GP can be as high as 67mg g à1DM in red GP (var.Cencibel)and as low as 1.68mg g à1in white GP (var.

Thompson,seedless)(Pe

rez-Jime nez et al.,2009).Some food processing methods such as extrusion increased extractable and bioavailable polyphenols (catechin and epicatechin)and reduced the NEP by reducing the degree of polymerisation (Khanal et al.,2009).Similar results were observed for the extractabil-ity of polyphenols in roasted peanut and almond skins (Yu et al.,2005,2006,2007;Garrido et al.(2008).These studies suggest that thermal processing may increase the extractability and bioavailability of some polyphenols while destroying heat sensitive polyphe-nols,in grape skin and seeds.Biological properties of GP polyphenols

Numerous studies have demonstrated that grape seed phenolics,particularly procyanidins,have many health bene?ts such as antimutagenic and anticarcinogenic activity (Joshi et al.,2000;Carini et al.,2000;Manten-a &Katiyar,2006;Yeh &Yen,2006;Bagchi et al.,2000;de Rezende,et al.,2009),antioxidant and anti-in?ammatory activities (Prior &Gu,2005;Sartor et al.,2002),prevention and delay of cardiovascular diseases (Brito et al.,2002;Vigna et al.,2003;Bagchi et al.,2003;Karthikeyan et al.,2009;Bradamante et al.,2004;Sano et al.,2005),increase in lifespan and retarded the onset of age-related markers (Valenzano et al.,2006).Some recent studies have also shown that taking grape seed extract (GSE)reduced food intake in rats and energy intake in human (Vogels &Plant-enga,2004).The main bioactive compounds responsi-ble for many reported health bene?ts of wine and wine by-products consist of antioxidant phenolics such as phenolic acids,anthocyanins,procyanidins and revera-trols (Shrikhande,2000;Yilmaz &Toledo,2004;).Antimutagenic and anticarcinogenic properties

A study by Bagchi et al.(2000)demonstrated that grape seed procyanidin extract (GSPE)was highly bio-available and provided signi?cantly greater protection against free radicals and free radical induced lipid oxi-dation and DNA damage than vitamins C,E or b -car-otene.Cytotoxicity of GSPE towards human breast,lung,gastric adenocarcinoma cells,while enhancing the growth and viability of gastric mucosal cells,was also observed in the same study.GSPE also exhibited protection against skin cancer by inhibiting UV-radia-tion-induced oxidative stress and activation of mito-gen-activated protein kinase and NF-j

B signalling in human epidermal keratinocytes (Bagchi et al.,2000;Mantena &Katiyar,2006;White et al.,2006).The study of Kaur et al.(2008)found that irrespective of source,GSE strongly inhibits LoVo,HT29and

SW480cell growth,with a G1arrest in LoVo and HT29cells,but an S and/or G2/M arrest in SW480cell cycle progression.GSE also induced Cip/p21levels in all three cell lines.Furthermore,an induction of apoptosis was observed in all three cell lines by GSE.These ?ndings suggest that GSE could be an e?ective alternative and complementary medicine against colo-rectal cancer because of its strong growth inhibitory and apoptosis-inducing e?ects.

It also has been reported that phenolics from grape seeds and skin inhibit some matrix proteases,such as leucocyte elastase and gelatinases,associated with in?ammation and cancer invasion (Sartor et al.,2002).The cell cultural study of Mertens-Talcott et al.(2008)show that the polypenol extracts from both red Musca-dine and Cabernet Sauvignon wine signi?cantly inhib-ited the growth of MOLT-4leukaemia cells.Wine extracts reduced cell viability up to 68%and cell num-bers up to 50%after 48h with muscadine extracts being more e?ective than cabernet sauvignon.These extracts also induced caspase-3activity and cell cycle arrest in the G2/M phase.The roles of anthocyanins in cancer prevention were extensively reviewed by Wang &Stoner (2008)and will not be repeated in this review.Grape seeds are rich in B type procyanidins.Many studies revealed that GSPE can serve as potential ther-apeutic agent for di?erent cancers.An in vitro study found that grape seed procyanidins inhibited pancre-atic carcinoma cells MIA PaCa-2and BxPC-3prolifer-ation in a dose-dependent manner and induced G1-phase arrest of the cell cycle in BxPC-3or mito-chondria-mediated apoptosis in MIA PaCa-2.Grape seed procyanidin also inhibited the adhesion and inva-sion potential of both cell lines in a dose-dependent manner through downregulation of MMP-2or MMP-9in pancreatic carcinoma cells (Chung et al.,2012).Grape seed proanthocyanidins also induced apoptosis of non-small cell lung cancer (NSCLC)cells,A549and H1299,in vitro through increased expression of pro-apoptotic protein Bax,decreased expression of antia-poptotic proteins Bcl2and Bcl-xl,disruption of mitochondrial membrane potential and activation of caspases 9,3and poly (ADP-ribose)polymerase (PARP).Further,administration of 50,100or 200mg GSPs kg à1body weight of mice by oral gavage (5days week à1)markedly inhibited the growth of s.c.A549and H1299lung tumour xenografts in athymic nude mice,which was associated with the induction of apoptotic cell death,increased expression of Bax,reduced expression of antiapoptotic proteins and acti-vation of caspase-3in tumour xenograft cells (Singh et al.,2011).Grape seed catechin and procyanidin B 4pretreatment was found to protect cardiomyocytes against doxorubicin-induced toxicity by decreasing reactive oxygen species generation as well as the num-ber of apoptotic cells,preventing DNA fragmentation,

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regulating the expression levels of the pro-apoptotic protein Bax-a and the antiapoptotic protein Bcl-2,and inhibiting apoptotic signalling pathways(Du&Lou, 2008).

There is growing evidence that resveratrol can pre-vent or delay the onset of various cancers,heart dis-eases,ischaemic and chemically induced injuries, pathological in?ammation and viral infections.As a chemoprevention agent,resveratrol has been shown to inhibit tumour initiation,promotion and progression (Jang et al.,1997).The review of Shanker et al.(2007) summarises the molecular mechanisms of resveratrol and its clinical bene?ts for human diseases.Resvera-trol induces apoptosis by upregulating the expression of Bax,Bak,PUMA,Noxa,Bim,p53,TRAIL, TRAIL-R1/DR4and TRAIL-R2/DR5and simulta-neously downregulating the expression of Bcl-2,Bcl-XL, Mcl-1and survivin.Resveratrol also potentiates the apoptotic e?ects of cytokines,chemotherapeutic agents and gamma-radiation.Pharmacokinetic and pharmaco-dynamic studies demonstrate that the main target organs of resveratrol are liver and kidney,and it is metabolised by hydroxylation,glucuronidation,sulf-ation and hydrogenation(Bishayee et al.,2010).Resve-ratrol(~25l M)potentiated GSE(35l g mLà1) induced colon cancer cell apoptosis via the activation of p53-dependent pathways(Radhakrishnan et al., 2011).This discovery suggests the importance of under-standing the potentiating e?ects of phytonutrients in combination as they would occur in nature rather than individually.

The cancer prevention mechanism of food polyphe-nols has been extensively studied.Many potential chemopreventive polyphenols may interrupt or reverse the carcinogenesis process by acting on intracellular signalling network molecules involved in the initiation and/or promotion of cancer(Manson,2003;Surh, 2003).The programmed cell death is considered one of the important targets in a preventive approach against cancer.Reversing the conversion of a normal cell to a malignant one is a complex process that involves active participation of a?ected cells in a self-destruc-tion cascade.In addition to signal transduction and regulation of proliferation and immune response,die-tary polyphenols readily interact with reactive oxygen species or free radicals to form relatively stable com-pounds,thus prevent cells from oxidative damage and onset of cancer.The major chemoprevention and che-motherapy mechanisms of dietary polyphenols may include(i)alteration of phase-I and phase-II drug-met-abolising enzymes,(ii)antioxidant properties,(iii) inhibition of protein kinases,(iv)blocking of receptor-mediated functions,(v)attenuation of protease activities,(vi)alteration of cell cycle checkpoint controls,transcription factor expression and apoptosis, (vii)inhibition of angiogenesis,invasion and metastasis,and(viii)epigenetic changes in promoter methylation and chromatin remodelling(Dashwood,2007;Kundu &Surh,2008).

Prevention of cardiovascular diseases

Epidemiological studies suggest that consumption of wine,grape products and other foods containing polyphenols is associated with decreased risk of car-diovascular disease.Cardiovascular disease is associ-ated with modi?cations in fatty acid metabolism and excessive lipid peroxidation of LDL.These oxidation products are also implicated in the formation of thromboxane,which leads?rst to enhanced platelet aggregation,then to artery blockage and?nally to thrombosis.The accumulation of lipid oxidation prod-ucts from LDL can be attributed to the low levels of plasma antioxidants.

A rat study showed that a15%GP in cholesterol diet(0.3%)produced a signi?cant reduction in choles-terol and triacylglycerols in the liver and serum.The diet contains15%GP reduced VLDL and LDL by50 and60–70%,respectively,while increase HDL level by 26%(Bobek,1999).Grape seed polyphenols reduce the risk of heart disease by inhibiting the oxidation of LDL(Shi et al.,2003a,b).Intravenous and oral administration of grape seed procyanidins was found to signi?cantly inhibit laser-induced thrombus forma-tion in the carotid artery of mice(Sano et al.,2005). Protection against myocardial ischaemia-reperfusion and myocardial injury in rats was reported by Bagchi et al.(2000)and Karthikeyan et al.(2009).GSEs rich in polyphenols exhibited higher e?ectiveness in reduc-tion in platelet adhesion,aggregation and generation of superoxide anion than pure resveratrol(Olas et al., 2008).Experimental studies indicate that grape polyphenols could reduce atherosclerosis by a number of mechanisms,including inhibition of oxidation of LDL and other favourable e?ects on cellular redox state,improvement in endothelial function,lowering blood pressure,inhibition of platelet aggregation, reducing in?ammation and activating novel proteins that prevent cell senescence(Dohadwala&Vita, 2009).

Procyanidins from GP inhibit human endothelial NADPH oxidase,the enzyme responsible for the increased production of reactive oxygen species, regardless of their polymerisation degree and galloyla-tion percentage.The procyanidin fractions even blocked NADPH oxidase activity in intact HUVEC, inhibiting ROS production at both extra-and intracel-lular levels.Therefore,grape procyanidins are suitable NADPH oxidase inhibitors,which could serve as models for therapeutic alternatives for cardiovascular diseases(A lvarez et al.,2012).Supplementation with the high dose of mixture of cathechin,ca?eic acid and resveratrol signi?cantly reduced the presence of

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atherosclerotic plaque by 40and 36%in the aortic sinus and in the ascending aorta,respectively Norata et al.(2007).Resveratrol alone also showed signi?cant antiatherogenic and anti-in?ammatory e?ects in an animal model of rabbits fed a hypercholesterolemic diet (1%cholesterol)Matos et al.(2012).CVD pre-ventative activities of anthocyanins,including results from in vitro cell culture and in vivo animal model systems as related to their multiple proposed mecha-nisms of action,were reviewed by Wallace (2011)and will not be repeated in this review.Antilipogenic properties

Vogels &Plantenga (2004)reported that GSE reduced food intake in rats and energy intake in humans.In an in vitro study by Moreno et al.(2003),GSE was found to signi?cantly inhibit pancreatic lipase,lipoprotein lipase and hormone sensitive lipase that responsible for the fat digestion and metabolism in human body.A study with rats fed high-fat diet shows that GSPE recti?es dyslipidemia associated with a high-fat diet in rats and repress genes controlling lipogenesis and VLDL assembling in liver (Baiges et al.,2010).Low-dose (25mg per kg body weight per day)GSPE treat-ment of high-fat-diet (HFD)fed rats signi?cantly reduced the adiposity index and the weight of all the white adipose tissue depots and reversed the increase in plasma phospholipids induced by the HFD feeding (Caimari et al.,2012).Chronic consumption of grape phenolics has shown to reduce obesity development and related metabolic pathways including adipokine

secretion and oxidative stress in a rat model (De

corde et al.,2009).These studies suggest that grape seed polyphenols may play a role in body-weight management.Antiageing activities

Grape seed extract has been shown to have a modula-tory role on age-related oxidative DNA damage and lipid peroxidation in central nervous system of rats (Feng et al.,2005;Balu et al.,2006).Aged rats given GSE showed improved memory performance,reduced reactive oxygen species production,decreased protein carbonyl level,increased thiol level and reduced hyp-oxic ischaemic brain injury in their central nervous systems.Recent studies show that resveratrol could exert neuroprotection against ischaemia,seizure and neurodegenerative diseases (Markus &Morris,2008).In the study conducted by Zhang et al.(2010),rat primary midbrain neuron-glia cultures were used to elucidate the molecular mechanisms underlying resve-ratrol-mediated neuroprotection.The results clearly show that resveratrol protected dopamine neurons against lipopolysaccharide (LPS)-induced neurotoxicity in concentration-and time-dependent manners through the inhibition of microglial activation and the

subsequent reduction in proin?ammatory factor release.Mechanistically,resveratrol-mediated neuroprotection was attributed to the inhibition of NADPH oxidase.Antioxidant activities of GP polyphenols in foods

Antioxidant activity is the most notable bioactivity of phenolic compounds from GP.The antioxidative characteristics have been widely studied,including scavenging of free radicals,inhibition of lipid oxida-tion,reduction in hydroperoxide formation and so on (Xia et al.,2010).Polyphenols are proven to be strong antioxidants against lipid oxidation in food system.Grape seed phenolics (GSE)were reported to inhibit lipid oxidation and warmed-over ?avour development in meat products such as raw and cooked beef products (Ahn et al.,2002,2007;Mielnik et al.,2006;Brannan &Mah,2007).GSE has reduced rancid ?avour development and associated primary and secondary lipid oxidation products in various meat products like raw and cooked beef,pork patties,turkey,?sh oil,frozen ?sh and ground chicken breast and thigh meat (Ahn et al.,2002;Banon et al.,2007;Brannan &Mah,2007;Brannan,2009;Carpenter et al.,2007;Lau &King,2003;Miel-nik et al.,2006;Pazos et al.,2005)without altering the pH,water activity,binding strength or yield of the meat.The minimum concentration level of GSE required to produce an antioxidant e?ect was 400l g g à1in cooked pork and 0.1%(w/w)in ground chicken (Lau &King,2003).However,GSE did alter the colour of both raw and precooked chicken breast patties (Brannan,2009).The degree of discoloration caused by adding GSE may depend on the variety of grape,type of meat and dosage.GSE may be more suitable for red meat than white meat.The GSE that contains less anthocyanins may be bet-ter choice as antioxidant in meat products.Antimicrobial activities of polyphenols from GP

In addition to antioxidant activities and therapeutic functions,many plant phenolic extracts have been shown to have antimicrobial activity against speci?c strains of bacteria such as Streptococcus Mutans ,Staphylococcus aureus,Escherichia coli and Candida albicans (O’Keefe &Wang 2006;Daglia et al.,2007),S.aureus,E.coli and C.albicans (C.albicans )(Papad-opoulou et al.,2005).Ahn and others (Ahn et al.,2007)reported that 1.0%GSE in cooked ground beef reduced the growth of E.coli O157:H7,L.monocytoge-nes,S.Typhimurim and A.Hydrophila by 1.5,2,1and 3LogCFU,respectively,during a 9-day storage.By using scanning and transmission electron microscopy,these authors also found grape seed polyphenols functioned as bactericidal,which caused disruption of

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the bacterial cell wall.It was reported that activities of GP extract against both spoilage and pathogenic bacte-ria including Aeromonas hydrophila,Bacillus cereus, Enterobacter aerogenes,Enterococcus faecalis, E.coli, E.coli O157:H7,Mycobacterium smegmatis,Proteus vulgaris,Pseudomonas aeruginosa,Pseudomonas?uores-cens,Salmonella enteritidis,Salmonella typhimurium, S.aureus and Yersinia enterocolitica(O zkan et al., 2004;Sagdic et al.,2011).The crude extract of GP at 10%inhibited the growth of the foodborne pathogens including Enterobacteriaceae and coliform bacteria, Salmonella,S.aureus in beef patties during the storage periods.The spoilage microorganisms including yeasts and moulds and lipolytic bacteria were also inhibited by5%of GP extracts in beef patties(Sagdic et al., 2011).These studies also indicate that the antimicrobial activity of crude GP extract in meat products is low. Adding5–10%of crude GP extract may cause many undesirable e?ects to the beef patty,such as reduced sensory quality and reduced protein digestibility. Resveratrol,which is rich in grape skin,has been reported to have strong antifungal and antibacterial activities.Fungi reported to be sensitive to resveratrol include human pathogens(C.albicans,Saccharomyces cerevisiae and Trichosporon beigelii)and plant patho-gens(Phytophthora palmivora,P.capisci,Aspergillus ?avus,Fusarium spp.and Verticillium spp.(Jung et al., 2005).The study of Paulo et al.(2010)veri?ed the antibacterial activity of resveratrol against Gram-positive bacteria.This study also used microscopic analysis and?ow cytometry techniques to reveal that the antibacterial e?ects of resveratrol were attributed to bacteriostatic action.The addition of resveratrol has allowed the identi?cation of changes in cell mor-phology and DNA contents.This suggests that the cell cycle is a?ected by resveratrol.

Therefore,polyphenols extracted from GP have the potential to be used for food preservation and medici-nal purpose to suppress the growth of pathogenic bac-teria and prevent oxidation of lipids.However,higher polyphenol concentration is required to achieve desired antibacterial activity against pathogenic bacteria com-pared to conventional antibiotics/antimicrobials. Thermal stability of polyphenols from GP

The bioactivity of polyphenols from di?erent plant sources are usually determined using compounds extracted at room temperature or refrigeration tem-perature and dried using freeze-drying technology to preserve their activity.Food processing conditions such as baking,steaming and extrusion are usually very harsh.Under such conditions,many bioactive compounds may undergo chemical degradation,isom-erisation or polymerisation and lose their activities. The most extensively studied stability of phenolics includes the enzymatic oxidation of fruit and vegeta-ble polyphenols during harvest,storage,transporta-tion and other handling.The thermal oxidation/ degradation of polyphenols during food processing such juice making,nut roasting and raw material dry-ing was also the subject of numerous studies.Both polyphenol content and antioxidant activity of foods decrease because of thermal processing and long-term storage(Klimczak et al.,2007;Hager et al.,2008). The study of Spanos et al.(1990)and van der Sluis et al.(2005)found that contents of polyphenols such as cinnamics,phloretin glycosides,procyanidins and quercetin derivatives in apple juice decreased during room temperature storage of apple juice.Drying of grape seeds at100and140°C resulted in18.6and 32.6%reduction in extractable total polyphenols, respectively,and reduced antioxidant activity of grape seeds compared to freeze-drying(Larrauri et al.,1997). The addition of lecithin in a solution containing tea catechins signi?cantly reduced the oxidative degrada-tion of the catechins at acidic pH and room tempera-ture(Lin et al.,2007).Heating decreased procyanidin and anthocyanin concentrations in freeze-dried GP signi?cantly(P<0.05).Reduction in procyanidin occurred when heated at60°C or above for8h with no further reduction when heating temperature increased from105to125°C.Total anthocyanin loss was highest at125°C(70%).No signi?cant loss of both procyanidin and anthocyaninin was observed when heated at40°C for up to3days(Khanal et al., 2010).When compared to freeze-drying,vacuum-dry-ing of fresh GP at60°C for24h also caused signi?-cant loss of total polyphenol,?avonoid and anthocyanin(Yu et al.,2011).

Controversial results were reported by Kim et al. (2006)who found that phenolic extracts of thermal processed whole and powdered grape seeds had higher in vitro antioxidant activity than those of untreated grape seeds.They also found newly formed low-molecular-weight phenolics in the extracts of heat treated grape seeds.Whether the newly formed phenolics were responsible for the increased antioxidant activity of GSE needs to be investigated.The study of(Friedman &Ju rgens,2000)demonstrates that ca?eic,chlorogenic and gallic acids are not stable to high pH and that the pH-and time-dependent spectral transformations are not reversible.By contrast,chlorogenic acid is stable to acid pH,to heat and to storage when added to apple juice.(à)-Catechin,(à)-epigallocatechin,ferulic acid,rutin and trans-cinnamic acid resisted major pH-induced degradation.Factors a?ecting the stability of anthocyanins are well known with pH being the most important factor for the colour of anthocyanins (Mazza&Brouillard,1987;.More information about the e?ects of non-thermal and thermal processing on anthocyanin stability in foods can be found in the

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reviews of Tiwari et al.(2009)and Patras et al.(2010),respectively,and will not be repeated in this article.Polyphenol –protein interaction

In addition to thermal stability,the interaction between food ingredients plays a vital role in success-ful food product development.Grape seed procyani-dins interact strongly with proteins leading to rapid the formation of protein –tannin aggregates,and the binding increases with the degree of polymerisation and molecular weight of procyanidins (de Freitas &Mateus,2001).Present knowledge indicates that this interaction is a?ected by parameters of the protein (molecular size,hydrophobicity,structural ?exibility),the polyphenol (degree of polymerisation,extent of galloylation,structural ?exibility)and the environ-ment (temperature,pH,ionic strength,presence of organic solvents and presence of carbohydrates)(Carvalho et al.,2006).To date,the most intensively studied polyphenol-ingredient interaction is tannin –protein interaction using bovine serum protein (BSA)with a few studies using b -lactalbumin (Prigent et al.,2009).This type of interaction is claimed to be responsible for the astringent taste of polyphenol-rich fruits and vegetables (Payne et al.,2009),haze forma-tion in beverages (Siebert,1999)and reduced bio-availability of both food protein and polyphenols (Skrabanja et al.,2000;Papadopoulou &Frazier,2004).However,the interactions between polyphenols other than tannins and food ingredients have rarely been reported.

Furthermore,considering the protein nature of enzymes,the interaction of polyphenols with digestive enzymes may reduce enzyme activity,thus reducing the digestibility of other food components such as car-bohydrates,proteins and lipids.The inhibition of digestive enzymes such as lipase (Moreno et al.,2003),proteases (Gonc alves et al.,2007),as well as glucosid-ases (McDougall et al.,2005;Gonc alves et al.,2011),because of interaction with grape seed procyanidins has been reported.The inhibitory e?ect increased with increasing degree of polymerisation of the procyanidin fractions.The inhibition is also accompanied by the formation of insoluble aggregates detected by dynamic light scattering and nephelometry (Gonc alves et al.,2011).

However,research ?ndings on the impacts of pro-tein –polyphenol interaction on protein digestibility have been controversial.He et al.(2006)reported that tea polyphenols at concentration of 50ppm inhibited the activities of a -amylase,pepsin,trypsin and lipase in bu?er solutions by 61,32,38and 54%,respectively.In contrast,other investigators reported that polyphenols,particularly procyanidins,have the ability to bind to dietary protein,thus

protecting it from rumen degradation,and increase protein availability in the small intestine of the host (Mueller-Harvey &McAllan,1992).A considerable interaction between polyphenols and proteins appeared during the hydrothermal treatment of buckwheat,and this interaction reduces the digestion of proteins through the small and large intestine.Microbial pro-cesses in the colon enhance the digestibility of pro-tein,blocked by polyphenols in hydrothermally processed buckwheat (Skrabanja et al.,2000).Adding carbohydrates into a tannin-BSA system induced a solubilisation of the protein/tannin complexes,with neutral and ionic polysaccharides displaying di?erent behaviours in this process.Pectin,xanthan,polygalac-turonic acid and gum arabic were much more e?ec-tive in solubilising the protein –tannin aggregates than glucose,dextran,b -cyclodextrin or arabinogalactan (de Freitas et al.,2003).Therefore,the presence of large amount of carbohydrate such as starch may reverse the enzyme –polyphenol interaction and protect enzyme activity.

Safety issue of polyphenol consumption

However,the pharmacological e?ects of these antioxi-dants are dose dependent and are a?ected by many factors including genotype of individuals.On the other hand,it has been reported that a number of antioxi-dants may have both anticarcinogenic and carcino-genic e?ects.Some of powerful antioxidants such as vitamin A,vitamin E,quecertin,catechins,procyani-din B2were reported to cause oxidative damage to cel-lar and isolated DNA (Sakano et al.,2005).It was also reported that feeding Swiss Webster mice with green tea extract epigalocatechin gallate at the dose of 50mg kg à1body weight caused liver necrosis and mortality in both male and female mice (Goodin et al.,2006).The review by Mennen et al.(2005)pointed out that certain polyphenols might have carcinogenic/genotoxic e?ects or may interfere with thyroid hor-mone biosynthesis;consumption of polyphenols may also inhibit non-haem iron absorption and may lead to iron depletion in populations with marginal iron stores;?nally,polyphenols may interact with certain pharmaceutical agents and enhance their biologic e?ects.

However,many adverse e?ects of polyphenols are dose related.Noticeable DNA damage was induced in mice spleen cells by incubating with higher concentration (150l M )of catechin (Fan &Lou,2008).Grape extracts were also found to promote mitomycin C (MMC)that induces sister chromatid exchange at concentration from 75to 300l g mL à1in human peripheral blood lymphocytes (Stagos et al.,2007).Among the phenolic compounds tested,ca?eic acid,gallic acid and rutin hydrate enhanced

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MMC-induced clastogenicity,whereas ferulic acid, protocatechuic acid,(+)-catechin,(à)-epicatechin and trans-resveratrol had no e?ect at concentrations between5and100l M.A considerable amount of evidence is accumulating,which supports the hypoth-esis that high-dose polyphenols can mechanistically cause adverse e?ects through pro-oxidative action (Martin&Appel,2010).Feeding male lambs on diets containing5and10%dry GP signi?cantly improved their growth performance(P<0.01)com-pared to the other treatments(Bahrami et al.,2010). The inclusion of GSE at levels0.6, 1.8and 3.6g kgà1body weight in broiler chicks of1–42 days did not a?ect the performance and the relative liver and pancreas weights,but reduced relative intestinal length at21days of age,increase ileal digestibility of crude protein and increased relative spleen weight at21and42days of age,respectively (Brenes et al.,2010).Therefore,it is important to consider the doses at which these e?ects occur,in relation to the concentrations that naturally occur in the human diet.

Applications of GP polyphenols in food systems

Some polyphenols have long been used in food prod-ucts.For example,anthocyanins from grapes and berries are used as food colourants(Shahidi&Nac-zk,2004).Food products forti?ed with plant extracts containing polyphenols are beverages including water or tea-based drinks,dairy products such as yoghurt, and special formulations such as‘smoothies’(Buch-wald-Werner et al.,2009).Signi?cant e?ort has been made over past decade to explore the potential of using GP to produce functional food ingredients, such as natural antioxidants for nutrition forti?cation and food preservation,health promoting grape seed oil and dietary?bre.GSE and GP extract are granted Generally Recognized As Safe(GRAS)and can be used as colour additive in fruit juice and?a-voured beverage as antioxidants(FDA,2003).GP pomace has been reported to use in the baked food products such as bread to increase the antioxidant activity of the bread and inhibit lipid oxidation of raw and cooked chicken(Peng et al.,2009;Sa yago-Ayerdi et al.,2009).In the study of Peng et al. (2009),bread was forti?ed with GSE,and the in vitro antioxidant activities,texture and colour of breads incorporating di?erent levels of grape see extract (300,600and1000mg per500g bread)were deter-mined to evaluate the e?ects of adding GSE on the quality of bread.Feeding chickens (3–6weeks old)GP extract at levels of0,30and 60mg kgà1body weight for3weeks did not a?ect chicken’s growth performance,but signi?cantly inhib-ited lipid oxidation of raw and cooked breast chicken patties compared with samples obtained from birds fed the control diet at20days and long-term frozen storage(6months)(Sa yago-Ayerdi et al.,2009). These results indicated that dietary GP polyphenols could be e?ective in inhibiting lipid oxidation of chilled and long-term frozen stored chicken patties. However,the production of puri?ed polyphenol extract is usually costly,and organic solvents such as ethanol,ethyl acetate and acetone are usually used(Masquelier,1987;Karvela et al.,2009).The use of organic solvent not only has harmful health impact on workers,but also generates new environ-mental problems.Although some e?orts have been made to avoid using organic solvents,the process for extraction,puri?cation and concentration of polyphenols is very tedious and costly(Shrikhande et al.,2000;Ochiai&Ueda,2007).Therefore,direct inclusion of GP in some food formulas,where GP function as polyphenol carrier,may be a better way for the polyphenol/antioxidant forti?cation of foods. Addition of0,5,7.5and10%deseeded GP in coo-kie formula increased dietary?bre and ashes sub-stantially in the cookies and did not a?ect the acceptability of cookies,although deseeded GP addi-tion imparted a darker colour to the cookies.How-ever,the higher the deseeded GP addition,the lower the net protein ratio,apparent digestibility and true digestibility of cookies(Canett Romero et al.,2004). This is most likely owing to the inhibiting e?ect of GP polyphenol on digestive enzymes.Incorporation of0.5–5%of grape seed?our in frankfurters led to a decline in the oxidation level of the products.The increment of grape seed?our in the frankfurters enhanced the protein,total dietary?bre and water-holding capacity of the treatments(P<0.05),but the colour values(L*,a*and b*)of frankfurters generally decreased(P<0.05)with increasing amount of grape seed?our(O zvural&Vural,2011). The possibility of using GP as feed was also stud-ied.Basalan et al.(2011)investigated the nutrient contents and in vitro digestibility of twenty-eight fresh GP samples from white and red wine grape varieties in Turkey.They found that in vitro disap-pearance of DM and NDF at48h determined using ruminal?uid was similar for pomace from both white and red grapes,but the in vitro disappearance of skin and seed DM was higher than that of stalk. This study suggests that fresh GP rich in skin and seed should be a suitable feed for ruminants and to non-ruminants with extensive cecal fermentation. However,inclusion of GP in the sheep diet to55% reduced digestibility of protein signi?cantly(Bau-mga rtel et al.,2007).GP may also be used to syn-thesise exo-polygalacturonase(exo-PG),pectinase, xylanase and cellulase by solid-state or submerged fermentation(D?az et al.,2012).

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Grape seed oil

The oil content of grape seeds was reported in range of 11.6–19.6%depending on the variety and maturity of grapes (Rao,1994;Llobera &Can ellas,2007).The fatty acid composition of grape seed oil also variety and maturity dependent.Major fatty acids of grape seed oil are linoleic (66.76–73.61%)acid,oleic acid (17.8–26.5),palmitic acid (6.35–7.93%)and stearic acid (3.64–5.26%),respectively (Beveridge et al.,2005;Rubio et al.,2009).It was found that polyunsaturated fatty acid (PUSFA)of oils from Cabernet Sauvignon and Royal Rouge pomace ranged from 60.9to 64.4%with high ratios of PUSFA/SFA (ranging from 2.80to 3.11)and high ratios of n à6/n à3(20.8–36.9)(Yi et al.,2009).The total unsaturated fatty acid accounts for more than 86%of the oil,and they were all essential fatty acids (EFA)(Baydar &Akkurt,2001).Dietary EFA was reported to determine the ?uidity of neuro-nal membrane and control the physiological functions of the brain Yehuda et al.(2005).EFA de?ciency dur-ing infancy delays brain development and in ageing accelerates the deterioration of brain functions.One study with human subjects found that high-monoun-saturated fatty acid diets lowered total cholesterol by 10%and low-density lipoprotein (LDL)cholesterol by 14%,while the high-density lipoprotein (HDL)remained unchanged (Kris-Etherton et al.,1999).A study involving a large number of nurses revealed that the group with lowest intake of linoleic acid exhibited the highest incidence of breast cancer (Eynard &Lopez,2003).

The antioxidant and fatty acid compositions of grape seed oil and thus its nutritional and cosmetic properties may be signi?cantly a?ected by the grape variety,growing conditions,oil extraction methods and degree of re?ning.Many researchers investigated the possibility of using supercritical CO 2?uid extrac-tion method to produce high-quality grape seed oil (Passos et al.,2009).The study by dos Santos Freitas et al.(2008)found that the most proper solvent for grape seed oil extraction is propane because oil sam-ples extracted with propane present a smaller amount of free fatty acids in the oil than samples extracted with carbon dioxide.However,owing to the high cost of supercritical ?uid extraction,commercial grape seed oil is mainly produced by traditional oil extraction methods such as hydraulic press and solvent extrac-tion.

In addition to phenolic antioxidants,grape seeds also contain non-phenolic antioxidants such as toc-opherols and b -carotene,both vitamins are potent antioxidants and are critical to human health.Toc-opherols and b -carotene are mainly concentrated in grape seed oil (Baydar &Akkurt,2001;Bravi et al.,2007).The content of tocopherols in grape seed oil

ranges from 265to 454mg kg à1depending on the extraction method,grape variety,growing location and growing conditions (Baydar &Akkurt,2001).a -Tocopherol was the most abundant tocopherol in the oil extracts,and c -and d -tocopherols were found with low concentrations,while b -tocopherol was not detected in the oil extracts (Baydar et al.,2007).The tocopherol and tocotrienol contents of grape seeds from 14di?erent varieties grown in Korea were deter-mined by Wie et al.(2009)using saponi?cation extrac-tion followed by normal-phase liquid chromatography,and it was found that the total concentration of tocopherol (T)and tocotrienol (T3)was in the range of 4.8–9.9mg 100g à1seeds (or 35.3–68.8mg 100g à1oil basis).The Muscat Bailey A cultivar had the high-est total tocopherol and tocotrienol contents,followed by Canner and Naples.c -T3ranged from 1.6to 4.9mg 100g à1seed (11.2–53.81mg 100g à1oil basis)and was the main isomer,followed by a -T3in most of the samples.

Grape seeds also contain certain amount of phytos-terols.Phytosterols are well known to contribute anti-arteriosclerotic activity.These sterols are concentrated in the grape seed oil.The total sterol content of grape seeds was reported to be 18530mg per kilogram oil.Among them,b-sitosterol is the most abundant (69.80–61.54%)followed by stigmasterol (11.87–16.03%),campesterol (10.79–9.28%),and sitostanol (3.47–3.97%)(Rubio et al.,2009).The concentration of each sterol changes with the variety and the maturity of grapes (Beveridge et al.,2005).The relatively high phytosterol content may make an important contribution to the health bene?t of grape seed oil.

Grape seed oil is a preferred cosmetic ingredient for damaged and stressed skin tissues.The regenerative and restructuring qualities of grape seed oil are most likely owing to its high antioxidant and sterol contents that may make it an attractive product for direct food consumption and skin care.

Grape seed protein

Relatives fewer studies were found for grape seed pro-tein.GPs/seeds are not considered as an important protein source as legumes and nuts,although grape seeds contain 11–13%proteins (Fantozzi,1981;Rao,1994;Gon i et al.,2005).The total protein content and the amino acid composition of grape seed protein may vary signi?cantly depending on the variety of grape,location and fertilisation conditions.Amino acid anal-yses of grape seed protein revealed high levels of essen-tial amino acids,but glycine,glutamic acid and aspartic acid were the most abundant amino acids found in GSP (Rao,1994;Zhou et al.,2010).The major protein component in grape seed protein isolate was found to be a globulin-link protein with subunit

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molecular weights varying from25.5to40.0kDa as determined by SDS-PAGE;the isoelectric pH of grape seed protein was found to be at the acidic pH of around3.8;grape seed protein possesses better solubil-ity,emulsifying capacity and emulsion stability than soy protein isolate,although its foaming capacity is poorer(Zhou et al.,2011).However,grape seed pro-tein was considered as non-digestible or resistant pro-tein(Saura-Calixto et al.,1991).Inclusion of GP in the sheep diet to55%signi?cantly reduced digestibility of protein(Baumga rtel et al.,2007).This is most likely attributed to the strong interaction between protein and tannins.The complexation between protein and tannin limited the digestibility of grape seed protein because tannin is believed to be an inhibitor of diges-tive enzymes(Gonc alves et al.,2007;Alipour&Rou-zbehan,2010).

The digestibility of puri?ed grape seed protein has not been reported.More research is needed to investi-gate whether procyanidins bound to grape seed protein can be released and absorbed in the gastrointestinal tract after eaten.

Dietary?bre in GP

Dietary?bres(DF)are de?ned by the Association of O?cial Analytical Chemists as‘the polysaccharides and remnants of plant materials that are resistant to hydrolysis(digestion)by human digestive enzymes’(Cho et al.,1997).DF includes many complex sub-stances,each having unique chemical structure and physical properties.Health bene?ts of dietary?bre are well documented(Slavin,2005;Anderson et al.,2009; Mann&Cummings,2009).

It is recognised that the physiological and physico-chemical e?ects of dietary?bres depend on the relative amount of individual?bre components,especially as regards to the soluble and insoluble fractions(Elleuch et al.,2011).Ideal?bre should include a balanced ratio of soluble and insoluble fractions(1:3)(Kunzek et al.,2002).Soluble?bres are characterised by their capacity to increase viscosity and to reduce the glyce-mic response and plasma cholesterol,protects against in?ammatory bowel diseases and to act as a prebiotic to improve host health(Anderson et al.,2009;Chawla &Patil,2010).Soluble?bre can be fermented in large intestine to produce short chain fatty acids that posi-tively a?ect major regulatory systems,such as blood glucose and lipid levels,the colonic environment and intestinal immune functions(Roy et al.,2006;Wong et al.,2006;Scholz-Ahrens et al.,2007).Insoluble ?bres are characterised by their high porosity,low density and ability to increase faecal bulk and decrease intestinal transit(Isken et al.,2010).The insoluble cer-eal?bre and whole grains were reported to be strongly associated with reduced diabetes risk in prospective cohort studies,indicating that other unknown mecha-nisms are likely to be involved(Isken et al.,2010). Dietary?bre is often intimately associated in the plant cell structure with other organic compounds,such as vitamins,phytochemicals,etc.,displaying their own biological activity(FAO,1998).GP is a?bre and poly-phenol-rich by-product of wine making.GP accounts for about20–25%of grapes crushed for wine making (Laufenberg et al.,2003)and contains up to75%of die-tary?bre and over60%of GP dry matter that was indi-gestible in vitro(Bravo&Saura-Calixto,1998).The indigestible GP dietary?bre includes pectin,cellulose, Klason lignin and polyphenols(Llobera&Can ellas, 2007;Gonza lez-Centeno et al.,2010;Deng et al.,2011). The composition of GP dietary?bre also depends on the variety of grapes and the part of pomace.The white GP had lower?bre concentrations(crude?bre, neutral detergent?bre and acid detergent?bre)than the red wine pomace(Baumga rtel et al.,2007).The dietary?bre of pomace and stem of Manto Negro red grape(V.vinifera)was studied by Llobera&Can ellas (2007).High percentage of soluble?bre(15%)in rela-tion to the total dietary?bre was found in the pomace, while high content of Klason lignin was found in both by-products,especially in the stem(31.6%).The Klason lignin fraction had important amounts of condensed tannins and resistant protein.Gonza lez-Centeno et al.(2010)investigated the dietary?bre compositions of pomace and stems from ten grape (V.vinifera L.)di?erent varieties(six red and four white).Both presented considerable quantities of DF, ranging from60to90%of total dry matter.The cell wall polysaccharides(CWP)composition of GPs and stems were very di?erent,pectic substances being the main component of the cell walls(40–54%of total CWP),while cellulose being the predominant CWP for the stems(40–49%of total CWP).Klason lignin accounted for around20–25%of DF in both GPs and stems.In addition,the pectin content and the degree of methyl-esteri?cation of uronic acids of GPs varied depending on the varieties of grapes.The study of Deng et al.(2011)with GP from two white wine grape varieties and three red wine grape varieties found that insoluble DF composed of Klason lignin(7.9–36.1% DM),neutral sugars(4.9–14.6%DM)and uronic acid (3.6–8.5%DM)weighed more than95.5%of total DF in all pomace samples from?ve grape varieties.White GP was signi?cantly lower in DF(17.3–28.0%DM) than those of red GPs(51.1–56.3%).Although data about the soluble and insoluble?bre contents of GP vary from study to study,there is no doubt that GP ?bre is low in solubility.Owing to the large quantity generated from worldwide wine and grape juice pro-duction every year,GP has potential to serve as an important source of insoluble?bre for functional food development.

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The results from a randomised,controlled parallel-group trial with thirty-four non-smoking adults show that consuming 7.5g day à1grape antioxidant dietary ?bre for 16weeks signi?cantly reduced total choles-terol (9%),low-density lipoprotein cholesterol (9%),and systolic and diastolic blood pressures (6and 5%,

respectively)(Pe

rez-Jime nez et al.,2008).The NEP bound to grape ?bre are not bioaccessible in the small intestine but can be partially released from ?bre matrix in large intestine by the action of gut bacteria (Selma et al.,2009;Saura-Calixto et al.,2010;Saura-Calixto,2011).A study with female Sprague –Dawley rats found that NEPA were partially depolymerised during their transit along the intestinal tract,as evidenced by the presence of (epi)catechin (EC)monomers and dimers in faeces and phase-II conjugates of EC in urine 24h after ingestion of NEPA-rich diet.More-over,NEPA were further metabolised by the intestinal microbiota into smaller metabolites including phenolic acids (Mateos-Mart?n et al.,2011).Therefore,GP ?bre could serve as a carrier for transportation of some polyphenols to the large intestine.Inclusion of GP in food products could result in the functional foods with bene?cial e?ects of dietary ?bre and grape polyphe-nols.

Conclusion

This review illustrates that the polyphenol composition of GP has been well characterised and their biological and functional properties are also intensively studied.The mechanisms of chemoprevention,anticardiovascu-lar disease and other disease prevention activities of grape polyphenols have been graduate revealed by researchers of di?erent disciplines all over the world.Therefore,GP has great potential to serve as a source of functional food ingredient.To optimise the health bene?ts and minimise possible negative health e?ects of GP,more studies are needed to set the proper dose of grape polyphenols,to characterise the properties of other GP components such as the oxidative stability of grape seed oil,solubility and weight management properties of GP ?bre and to evaluate the sensory quality and consumer acceptance of food products developed from GP or components of GP.

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葡萄的二次结果技术 在葡萄生产中，有时芽眼受到冻害或霜害以及植株生长过旺，一次果偏少达不到预定的产量时，可用二次果加以弥补。一般二次果粒小，果皮较厚，着色好，较耐贮运，但含糖量低。如完全成熟也能形成优质果。有些品种容易形成二次果，如玫瑰香、巨峰系品种，康拜尔系品种等；有些则不易形成二次果，如龙眼、牛奶等。它比一次果晚熟15-20天，中、早熟品种已迅，龙眼等晚熟品种未熟，正好弥补市场空缺，也能取得较高的经济效。人工刺激形成二次果的方法有以下几种 一、利用冬芽二次梢结果。在带不出果穗的机关报梢长出10-15片叶时，进行摘心。抑制主梢 生长并暂时保留摘心口下的2个夏芽副梢，延缓主梢上的冬芽萌发，促进花序分化，其余副梢一律抹去。待暂时保留下来的副梢半木质化后，再从基部剪去，刺激主梢上的冬芽萌发，结二次果。一般主梢摘心口下的第一个冬芽，结实力较弱，第第二、第三外冬芽结实力较强。从主梢摘心至逼冬芽带出花序，应在20-30天内为宜，以免影响二次果的成熟。 二、第三外冬芽结实力较强。从主梢摘心至逼冬芽带出花序，应在20-30天内为宜，以免影响二次果的成熟。

二、利用夏芽副梢结果。开花前，在夏芽未萌发的节上剪截，促发副梢，使其带穗。如不带穗，在副梢上留2-3个叶，再进行剪截，直至果穗长出为止。若对副梢摘心过早，新抽生的二次副梢多为发育枝，要经过2-3次连续摘心，即有花序出现。 三、在生长季节喷洒300-1000倍矮壮素，可减缓夏芽萌发和新梢生长速度，促进副梢抽生花序，增加二次果的产量。 二次结果，可以充分发挥葡萄的增产潜力，尤其在遭受自然灾害、花果不足的情况下，对弥补当年产量有特殊的意义。但二次结果的植株，消耗养分较多，在肥水供应不足的情况下，对弥补当年产量和花芽分化带来不良影响，必须在栽培管理水平较高的果园，才能进行。

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葡萄品种杂交育种技术

葡萄品种杂交育种技术 全世界有4万多个葡萄品种，这些品种是如何产生的呢？ 除了大自然优胜劣汰自然留下的品种外，绝大部分是人类作用的结果，主要有芽变育种，杂交育种，辐射育种，诱变育种，转基因等等，其中杂交育种的品种比例最大，栽培面积也最大。 下面就杂交育种技术加以介绍，（也是本人的本职工作），望大家不要见笑：）杂交简介： 杂交是指通过选择两个亲本通过花粉传递的方式，将其融合，使其后代性状互补，形成具有双亲优良性状的后代株系的常规育种方法。举例说明：“玫瑰香”品种是英国育种家以“白玫瑰香”为母本，“黑汉”为父本杂交育成，其母亲白玫瑰香为绿色，枝条成熟度不好，果实香气不足，很难大面积推广，而父亲黑汉果粒小，无香味，更无法大面积推广，双亲各取其优点，通过杂交，不但继承了母亲“白玫瑰香”的香味，还继承了父亲“黑汉”的枝条成熟度好，果实黑色，等优良性状，把黑汉和白玫瑰香的共同优点继承，而摒弃了其缺点，造就了一个闻名世界的品种“玫瑰香”。再如我院选育的“巨玫瑰”，母本为香味浓郁的玫瑰香四倍体芽变沈阳玫瑰，父本为巨峰，“巨玫瑰”即继承了母本的香味，也遗传了巨峰的大部分优良性状，是我国欧美杂交种选育的一大突破。 1 采集花粉 将用作父本的花序取下，用手掌揉碎，摊于纸片上阴干。24小时后将干燥的花粉混合物收集于自封袋中备用。

2 去雄 在开花前1-2天，用镊子小心翼翼的将每串花的每朵小花的花帽和花药摘掉，同时不能碰伤柱头和果肉。

摘除了花药的花序 3 授粉 用自封袋套住已去雄的果穗，自两端向中间封死，然后轻轻抖动自封袋，使花粉混合物飞溅起来。

用手轻轻抖动，使花粉混合物飞溅至柱头

金手指葡萄设施栽培技术要点

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作物转基因育种研究进展 摘要：近年来，植物基因工程取得了辉煌的成就，而转基因技术由于其巨大的产业价值,特别是在作物品质改良、产量和抗逆性提高等方面的明显优势,一直是国际农业高新技术竞争的焦点和热点。本文主以棉花、玉米、水稻为例就转基因育种技术在作物上的研究进展进行相关的介绍。 关键词：作物，棉花，玉米，水稻，转基因育种，研究进展 植物转基因技术是指利用重组技术、细胞DNA培养技术或种质系统转化技术将目的基因导入植物基因组，并能在后代中稳定遗传，同时赋予植物新的农艺性状，如抗虫、抗病、抗逆、高产、优质等。常规育种常常受有性杂交亲和性的制约，而利用转基因技术可以打破物种界限、克服有性杂交障碍，快速有效地创造遗传变异，培育新品种、创造新类型，大大缩短新品种育成的时间。因此，随着现代生物技术的迅速发展，植物转基因技术也蓬勃发展［1］。 1 转基因棉花育种的研究与进展 近年来，随着基因工程技术的不断发展，利用生物技术来创新棉花种质资源和培育新品种是一条非常有效的途径，极大地推动了棉花遗传育种的发展［2］。中棉所是世界上唯一可以同时采用农杆菌介导法、花粉管通道法、基因枪轰击法快速获得转基因抗虫棉新材料的技术平台,能将植物嫁接技术成功应用于转基因棉花的快速移栽,成活率超过90%。未来3~5年,中棉所将挖掘、整合与优化抗病、抗除草剂等基因10个,筛选高产因子、高品质纤维等基因或分子标记150个,创造转基因棉花育种新材料100份以上,培育重大新品种(组合)3~5个。 1.1转抗虫基因 1991年成功将外源Bt基因导人棉株中，1992年人工合成了全长1824bp的CrylAb和CrylAc融合的GFMCry1A基因，并于1993年采用农杆菌介导法和外源基因胚珠直接注射法成功导入晋棉7号、中棉12、泗棉3号等主栽品种，获得了高抗棉铃虫的转基因棉花株系；包含CryIAc和AP基因双价抗虫基因载体，通过农杆菌介导转化冀合321胚性愈伤组织，经6代筛选后培育出抗棉铃虫90%的纯合品系，且农艺性状均优于对照。 1.2转抗黄萎病相关基因 利用花粉管通道法和农杆菌介导转化法将菜豆中的几丁质酶和烟草中的葡聚糖酶基因转入棉花，并从转基因高世代材料中筛选出了高抗黄萎病的品系；将天麻抗真菌蛋白基因用花粉管通道法转化天然彩色棉主栽品种，从高世代系中选育出既抗枯萎病又抗黄萎病的兼抗材料；将葡萄糖氧化酶基因(GO)转入棉花，转基因后代对枯萎病和黄萎病抗性均有显著提高，部分材料抗性达到抗病水平。1.3转抗除草剂基因 1997年由美国孟山都公司推出抗除草剂棉花抗性品种，他们从土壤农杆菌变种CP4中分离到编码抗草甘膦酶的基因，并通过农杆菌介导法转化珂字棉312，把该基因导入棉花植株，从而使其对草甘膦产生抗性。采用中棉35下胚轴为材料，将草甘膦突变基因aroAM12导入到棉花中，获得65棵再生植株，通过Southern及Western试验验证了该基因的导入和表达状况，结果表明，转化株对草甘膦具有很高的抗性；将抗草甘膦基因aroAM12和抗虫基因Btslm一起整合到一个载体中，并以抗草甘膦基因作为选择标记，通过转化棉花品种石远321后获得了抗草甘膦和抗棉铃虫的再生株。

超级杂交水稻育种研究进展.

超级杂交水稻育种研究进展 （袁隆平） H前，中国人口冇13亿，人均对耕地仅1.4亩，预11-2030年人11将增至16亿，人均町耕地将减少到Im 左右。面对人口增长爪力和耕地减少的严峻形势，为保障粮仅安全，农业部于1996年启动实施了超级水稻育种计划。 技术路线 □ 育种实践表明，通过育种提高作物产量，可川纳出两条仃效途径：一是形态改13,二是杂种优势利用。单纯的形态改良，潜力仃限；杂种优势不与形态改良结合，效果较差。相关育种途径和技术，包括基因工程在内的W技术，彊终将落实到优良的形态和强人的杂种优势匕才会对提高产量冇贡献。但是, 疗种进一步向更高层次发展，也必须依靠卞物技术的进步。

形态改良 优良株型是高产的基础。口从1968年Dr.Donald提出理想株型概念肩，很多水稻育种家特别注总这一重要课题，并设想了多种高产水稻模型。其屮著名的是国际水稻研究所Dr.Khush提出的“新株型"稻，其主耍特征是：①大穗，每穗250粒；②分藥少, 侮株 3~4个有效分簾；③短而壮实的秆。这种模型是否高产，还冇待实践证明。发现超尚产品种仃如下形态特征。 A.高冠层 上-叶叶片应长、直、窄、凹、厚。长而直的叶子不仅叶面枳大而11能两而受光又耳不遮荫，因此能更右效地利用光能；窄叶所占的空间面积小，能增加有效的叶面积指数；凹字形可使叶片坚挺不披；厚叶光合功能强IL不易早哀。总Z,具仃这种形态特征的水稻品种，能冇最大的冇效叶面积指数和光合功能，为超高产提供充足的光合产物即冇机源。

B.矮穗层 成熟期稻穗顶部离地面仅60?70cm, 这种结构由于重心卜?降，可使植株高度抗倒 伏。抗倒是培育超高产水稻必备的特性。 C?中大穗_ □毎穗谷疏约6克，每亩16?17丿总。稻谷产吊「=生物学产htx收获指数。理论I：,英产吐潜力为1000公斤/ 市。现行的矮秆品和，收获指数(HI)L2很高(>0?5), 进一步处场收蕊指数L2札严1仃限，W此，上加应依如提髙生枷％产幕以进氏-提高稻谷产量。 □从形态学观点來看，提高植株高度是提高生物学产啟有效而町行的方法。然而这种方法会引起僧伏。为解决这个问题，不少冇种家正试图使茎秆更?壮，但此举会导致收获指数下降，因此，很难达到超局产。上述山叶片组成的高叶冠层植株模型能同时将高生物学产量、髙收我指数和高度抗倒伏二者较好的统一起来, 从而能实现题W产。

葡萄

巨峰葡萄一年两熟栽培技术 摘要：桂南巨峰葡萄一年两熟栽培技术增加一造产量，本文从催芽、抹芽定梢、摘心、合理疏花疏果、葡萄套袋技术及采收时间等方面详细介绍了巨峰葡萄一年两熟的栽培技术、水肥管理及病虫害的防治，为葡萄生产者进一步了解并应用该技术提供参考。葡萄是喜温的果树植物，从萌芽到果实成熟所需≥10℃的活动积温都不相同，研究表明，极早熟品种要求2100-2500℃，早熟品种2500-2900℃，中熟品种2900-3300℃，晚熟品种3300-3700℃，极晚熟的品种则要求3700℃以上的活动积温[1]。葡萄是果树中花芽较易形成的树种之一，葡萄花芽具一年多次分化习性，在桂南热量资源丰富，光照强，几乎无霜冻，只要在管理上加以特殊处理（如采用全部落叶修剪，破眠催芽）等，就能实现一年两次结果（夏采和冬果）。目前，我国南方葡萄生产大多为一年一熟。为此，很早就有学者开展了葡萄一年双收的栽培技术研究工作，并取得了较大的进展[2，3，4，5，6，7]。但基本上是在第一造果未采收前处理获取二次果，如果第一造挂果多，第二造就很难形成产量，而且这样的二造果成熟很不一致。本栽培摸式合理利用巨峰葡萄花芽分化容易的特点，一年内收获生育期完全不重叠的两造产量；不仅可提高单位面积产量，提高葡萄的品质，延长鲜果的供应期，调节鲜果市场的淡季供应，还能显著的提高经济效益。但是，二次结果增加树体养分的消耗，如果管理不当，将影响次年夏果的产量和品质，甚至造成次年严重减产。另外，南宁地处亚热带，高温多雨，对葡萄的优质栽培有很大的影响，雨水过多，易形成洪涝灾害，影响根系对营养物质的 吸收，还容易滋生病害，对葡萄的生长、成熟和浆果品质带来不利影响。因此，在一年中要获得高产、稳产、品质好的双季葡萄，需要精细的管理，要有与之配套的栽培管理技术。1.品种的选择供鲜食的两熟葡萄应选择果粒大、着色易、果粒整齐、品质好，抗逆性强、容易形成花芽、早结丰产的品种作主栽品种。本文以巨峰葡萄为例。 2.一年两熟栽培管理方法本文介绍的一年两熟栽培技术是指在气候条件合适和良好土肥水条件的基础上，在一造果采收后，经过一个月左右的恢复期，采用全部落叶修剪，催芽破眠，刺激冬芽当年萌发，开启另一个生长周朝，生产第二造果（冬果）。应用该技术进行生产时，两造果的配套管理十分重要。2.1第一季果的管理 2.1.1适时破眠催芽葡萄属于落叶果树，低温是葡萄通过休眠的必要条件，而桂南巨峰葡萄冬季休眠期间的低温远远满足不了其休眠所需的低温要求。若在休眠不足的状态下萌芽，往往会造成葡萄萌芽率低，萌芽开花不整齐等生理障碍。用催芽剂打破休眠，能提高萌芽率和促进萌芽整齐，并能提早萌芽，提早成熟。既可获得较高的经济效益，同时也可以保证生产二造果（冬果）时有足够的生长时间。在南宁可于2月上中旬催芽。具体方法是：视结果母枝健壮情况留6-10芽修剪，将修剪好的结果母枝用49%单氰胺20倍溶液（加适量胭脂红，方便检查是否漏涂）均匀涂抹在冬芽上，但顶端1-2个芽不处理。处理前后必须灌水。 2.1.2抹芽定梢分三次进行：第一次把不定芽、接果母枝基部1-2个弱芽抹掉；第二次将副芽和极强旺梢抹掉；第三次在坐稳果后尽快进行，将过密枝、徒长枝、坐果不好的枝条剪掉，并把枝条均匀绑好，保证架面通风透光；2.1.3摘心在葡萄初花期对一次果枝或营养枝摘心，一次果枝花序以上留8-10片叶，营养枝留10-12叶摘心，每次摘心后尽早抹除刚萌动的副梢，只留顶端副梢，顶端副梢留2-3叶反复摘心，营养枝顶端副梢3-5叶反复摘心。摘心可以暂时终止该枝的延长生长，减少梢枝幼叶对养分、水分的消耗，促进坐果，促进留下的叶片迅速增大并加强同化作用，使树体内的养分重新分配，对结果枝来说，还可以改善花序（或

葡萄标准化栽培技术

葡萄标准化栽培技术 哈密有着得天独厚的气候、水土条件，是生产优质绿色葡萄的产区。由于在本地生产的无核葡萄具有肉脆、含糖量高、品质好、既可鲜食又可制干等特点。所以，深受国内外客商及广大消费者的喜爱，产品远销国内外。为了让广大种植户掌握规范化的葡萄管理技术，进一步提高葡萄品质，下面，我们将为大家详细介绍葡萄种植的管理技术。 一、葡萄开墩出土和上架 葡萄开墩出土的温度指标为旬平均气温10～13℃，这里所说的旬平均气温就是指十天之内的平均气温，也就是杏树开花的时候。我区一般在4月10日前后葡萄开始开墩，4月20日之前开完。但为了预防晚霜冻害，具体时间要参考当时的天气预报。开的过早容易受到晚霜的危害，过晚会影响葡萄当年的生长。葡萄开墩出土和上架可采用以下技术规程： （一）葡萄开墩出土技术要点 葡萄开墩最好分两次进行，第一次先除去枝蔓外较厚的覆土，目的是有利于在葡萄发芽前快速出土。第二次是在确定近期完全没有晚霜冻害，再把枝蔓外剩余的覆土全部进行清理。开墩时应小心，尽量不要损坏葡萄枝蔓，以免造成伤流和影响产量。 （二）葡萄上架技术要点 葡萄出土后，枝蔓应该及时上架。目前，哈密地区大部分是采用棚架，所以，种植户应该把棚架的主蔓在架面上按40～50cm的距离均匀摆放，枝蔓间不应交错纠缠。摆放好葡萄枝蔓后，要在枝蔓的关键部位与棚架进行捆扎，以免风刮或自动弹回。捆扎采用活套捆扎技术，这样既能防止捆扎处产生缢痕而影响养料和水分的输送，也可避免因缢痕而发生的折断。捆扎材料可以选用布条、麻绳等。在葡萄开墩出土的同时，还要将墩沟清理平整，及时灌好开墩水，防止枝蔓抽干，促进新梢生长和花序的形成。 二、葡萄施催芽肥

木本植物多倍体育种研究进展

植物学通报 2005, 22 (3): 375￣382Chinese Bulletin of Botany 木本植物多倍体育种研究进展① １ 李　云 ２冯大领②1(北京林业大学生物科学与技术学院, 教育部林木花卉遗传育种重点实验室 北京 100083) 2(河北农业大学生命科学学院 保定 071001) 摘要 本文从木本植物多倍体育种资源、多倍体育种途径和多倍体鉴定等方面, 对国内外木本植物的多倍体育种研究进展进行了综述。与农作物相比, 尽管木本植物在多倍体育种中有其缺陷, 但木本植物丰富的物种资源及无性繁殖等特性较大程度地发挥了多倍体育种的优越性。 关键词 木本植物, 多倍体育种, 育种资源 Advances in Research into Polyploidy Breeding of Woody Plants 1 LI Yun 2FENG Da-Ling ②1(College of Biological Science and Biotechnology,Beijing Forestry University , Key Laboratory for Genetics and Breeding in Forest Trees and Ornamental Plants, Ministry of Education , Beijing 100083) 2(College of Life Sciences, Agriculture University of Hebei , Baoding 071001) Abstract Advances in polyploidy breeding of woody plants is reviewed according to several factors, including polyploidy breeding resources, approaches and identification. Compared with polyploidy breeding of crops, that of woody plants remains superior because of the abundant spe-cies resources and asexual propagation, though there are limitations. Key words Woody plant, Polyploidy breeding, Breeding resources ①国家自然科学基金项目(30471412)和北京市自然科学基金项目(6042020)资助。 ②通讯作者。Author for correspondence. E-mail: shuqfeng@https://www.wendangku.net/doc/ea12384708.html, 收稿日期: 2003-12-22 接受日期: 2004-04-12 责任编辑: 崔郁英, 白羽红 多倍体普遍存在于植物界(浙江农业大 学, 1980; 何启谦, 1992)。在被子植物中约有 1/3或更多的物种是多倍体, 裸子植物中多倍 体物种较少, 约有5%。染色体加倍是变异发 生的重要途径之一, 对物种的进化以及育种 都有很重要的意义。 多倍体育种源于20世纪初(蔡旭, 1988), 到了20世纪30年代, 多倍体育种逐渐广泛应 用于农作物品种的培育中, 尤其自1937年秋水仙碱对诱发植物多倍体的巨大效果被证实以后, 利用植物碱和异生长素等化学药剂诱导多倍体的研究日益增加, 并取得一定成果。在林业上, 树木多倍体育种成果最引人注目的是20世纪30年代以后发现了一系列的天然树木三倍体, 从此人工诱导树木多倍体的工作在许多木本植物上广泛展开, 探索出了许多适于诱导树木多倍体的方法。最早育成的树木多倍体是红花七叶树(南京林产