摘要:【目的】探索光質(zhì)條件改變對陽光玫瑰葡萄冬果品質(zhì)及糖類物質(zhì)代謝變化的影響,為提高葡萄冬季果實品質(zhì)提供參考依據(jù)?!痉椒ā吭诙l(fā)育階段對陽光玫瑰葡萄植株進行LED燈照射補光處理,分別為紅光、藍光和白光,以不補光處理為對照;分別在硬果期(E-L33)、果實膨大期(E-L35)、果實未完全成熟期(E-L37)和果實完全成熟期(E-L38)各采樣1次,對果實樣品的縱徑、單粒重,可溶性固形物、可滴定酸、果糖、葡萄糖和蔗糖含量,蔗糖合成酶(SS)、蔗糖磷酸合成酶(SPS)和中性轉(zhuǎn)化酶(NI)活性及其基因相對表達量進行測定。【結(jié)果】白光對果??v徑的伸長影響較大,在E-L38時期較對照顯著提高4.94%(P<0.05,下同)。紅光處理促進果實可溶性固形物含量增加,在E-L38時期較對照顯著提高11.26%。藍光處理對果實可滴定酸含量的降低有促進作用,在E-L37時期較對照顯著降低15.54%。在E-L37和E-L38時期紅光處理促進果實果糖含量增加,分別較對照顯著提高11.14%和22.93%。紅光處理對果實葡萄糖含量積累有促進作用,在E-L38時期較對照顯著提高17.51%。在E-L35和E-L37時期,紅光處理的SS活性分別較對照顯著提高18.50%和13.39%。在E-L38時期,紅光、藍光和白光處理均對果實SS和SPS活性有促進作用,紅光處理的NI活性較對照顯著降低17.71%。紅光處理使VvSS和VvNI基因相對表達量在E-L35和E-L38時期分別較對照降低10.54%、20.90%和11.57%、28.50%。果實成熟期,藍光處理的VvSPS基因相對表達量較對照顯著降低22.74%?!窘Y(jié)論】紅光顯著促進陽光玫瑰葡萄冬果的可溶性固形物、果糖和葡萄糖含量的積累,藍光有助于可滴定酸含量的降低。紅光、藍光和白光在果實成熟期對SS和SPS活性均有促進作用。可根據(jù)陽光玫瑰葡萄冬果對光照環(huán)境及果實品質(zhì)的需求選擇合適的人工光源進行補光。
關(guān)鍵詞:陽光玫瑰葡萄;補光處理;果實品質(zhì);糖代謝;酶活性
中圖分類號:S663.1文獻標(biāo)志碼:A文章編號:2095-1191(2024)08-2286-09
Response mechanism of Shine Muscat grape winter berries quality and sugar metabolism to light quality
SHI Xiao-fang LIN Ling HUANG Qiu-mi BAI Xian-jin BAI Yang3,HUANG Gui-yuan HAN Jia-yu LI Hong-yan CAO Xiong-jun1*,GUO Rong-rong1*
(1Grape and Wine Research Institute,Guangxi Academy of Agricultural Sciences,Nanning,Guangxi 530007,China;2Guangxi Crop Genetic Improvement and Biotechnology Laboratory,Nanning,Guangxi 530007,China;3GuangxiZhencheng Agricultural Co.,Ltd.,Nanning,Guangxi 530105,China)
Abstract:【Objective】Exploring the effects of changes in light quality conditions on the quality and sugar metabolism of winter berries in Shine Muscat grapes,providing reference for improving the quality of grape winter berries.【Method】During the winter berry development stage,Shine Muscat grape plants were treated with LED lights to supplement light,including red light,blue light,and white light.The control group received no supplementary light treatment.Sampleswere taken once each during the hard berry stage(E-L33),berry swelling stage(E-L35),berry immature stage(E-L37),and berry fully mature stage(E-L38).The longitudinal diameter,single berry weight,total soluble solids content,titratable acid content,fructose content,glucose content,sucrose content,sucrose synthase(SS)activity,sucrose phos-phate synthase(SPS)activity,neutral invertase(NI)activity,and relative gene expression levels of berry samples were measured.【Result】White light treatment had great effect on the elongation of berry longitudinal diameter,which was 4.94%significantly higher than the control(P<0.05,the same below).Red light treatment promoted an increase in soluble solids content in berries,with a significant increase of 11.26%compared to the control during the E-L38 period.Blue light treatment had promoting effect on the reduction of titratable acid content in berries,with a significant decrease of 15.54%compared to the control during the E-L37 period.During the E-L37 and E-L38 periods,red light treatment pro-moted an increase in fruit fructose content,with significant increases of 11.14%and 22.93%compared to the control re-spectively.Red light treatment had promoting effect on glucose accumulation berries,with a significant increase of 17.51%compared to the control during the E-L38 period.During the E-L35 and E-L37 periods,the SS activity treated with red light significantly increased by 18.50%and 13.39%compared to the control respectively.During the E-L38 pe-riod,red light,blue light,and white light treatments all had promoting effect on SS and SPS activities in berries,and the NI activity of red light treatment was significantly reduced by 17.71%compared to the control.Red light treatment re-duced the relative expression levels of VvSS and VvNI genes by 10.54%,20.90%,11.57%and 28.50%respectively com-pared to the control during the E-L35 and E-L38 periods.During the berry fully mature stage,the relative expression level of VvSPS gene treated with blue light decreased by 22.74%compared to that of the control.【Conclusion】Red light signifi-cantly promotes the accumulation of soluble solids,fructose,and glucose contents in the winter berries of Shine Muscat grape,blue light helps to reduce titratable acid content.Red light,blue light and white light all have promote effect on SS and SPS activities during berry ripening.Suitable artificial light sources for supplementary lighting can be chosen based on the requirements of sunlight environment and berry quality for Shine Muscat grape winter berries.
Key words:Shine Muscat grape;supplementary light treatment;berry quality;sugar metabolism;enzymatic acti-vity
Foundation items:National Natural Science Foundation of China(32360728);Guangxi Natural Science Foundation(2020GXNSFBA159019);Basic Research Project of Guangxi Academy of Agricultural Sciences(Guinongke 2023YM113,Guinongke 2021YT126)
0引言
【研究意義】光是影響植物形態(tài)發(fā)生的重要信號(de Wit et al.,2016),葡萄(Vitis vinifera L.)是喜光、長日照植物,光照時數(shù)、光照強度等顯著影響葡萄生長發(fā)育。由于避雨栽培和一年兩收栽培技術(shù)的推廣應(yīng)用,廣西葡萄產(chǎn)業(yè)發(fā)展迅猛,逐漸成為專家們認(rèn)可的葡萄特殊優(yōu)勢栽培區(qū)(白先進等,2008)。根據(jù)國家統(tǒng)計局?jǐn)?shù)據(jù)顯示,截至2022年,廣西葡萄栽培面積約3萬ha,居全國前十,鮮食葡萄是廣西單位面積種植效益最好的果樹,對廣西農(nóng)民增收和鄉(xiāng)村振興起了重要的推動作用。糖、酸及花色苷等果實品質(zhì)均受光條件的影響,廣西葡萄冬果成熟關(guān)鍵期出現(xiàn)較多陰雨天,光照強度和時長不足,以及光質(zhì)差易導(dǎo)致冬季葡萄出現(xiàn)果粒偏小、果實糖度低、酸度高等品質(zhì)差問題(李洪艷等,2016)。葡萄一年兩收栽培模式下,研究冬果品質(zhì)及糖代謝響應(yīng)光質(zhì)機理,改善酸高糖低的現(xiàn)象,對廣西葡萄產(chǎn)業(yè)發(fā)展具有重要意義。【前人研究進展】光照是影響果實成熟、風(fēng)味物質(zhì)代謝最重要的環(huán)境因素之一(Dieleman and Meinen,2007;侯曉健等,2022)。補光可促進葡萄芽的發(fā)育(王欣欣等,2009),提高可溶性固形物和花青素含量(Ovadia et al.,2013;Kondo et al.,2014)及果實品質(zhì)(Li et al.,2017;時曉芳等,2021a),促進香味形成(Friedel et al.,2016);藍光可提高夏黑葡萄超氧化物歧化酶活性,紅光可提高其過氧化物酶活性(余陽等,2015);適宜比例的紅藍光處理能增強赤霞珠和LN33幼苗的光合效率(魏云春等,2023)。糖類物質(zhì)是影響果實甜度、酸、風(fēng)味等品質(zhì)的關(guān)鍵因素,葡萄果實含糖量可達15%~28%,以葡萄糖和果糖為主,而蔗糖含量較少(房經(jīng)貴和劉崇懷,2014;李棟梅等,2022)。葡萄糖和果糖能顯著誘導(dǎo)葡萄花青素苷的積累和基因表達(Zheng et al.,2009)。與風(fēng)味品質(zhì)有關(guān)的各類糖中,以果糖甜度最大,葡萄糖風(fēng)味最佳,在可溶性固形物含量相近的情況下,果糖和葡萄糖含量高更利于甜味的增加,也是葡萄形成自身獨特風(fēng)味的重要原因之一(秦巧平等,2005;冷翔鵬等,2011)。轉(zhuǎn)化酶主要將蔗糖水解為葡萄糖和果糖,蔗糖磷酸Z6BXuIR6N+WVjOyF172oVIRY34L56FWa3oOHSBTePJg=合成酶(SPS)是一種可溶性酶,是調(diào)節(jié)蔗糖合成的關(guān)鍵酶之一,蔗糖合成酶(SS)活性分為分解方向活性和合成方向活性。劉林等(2008)研究不同反光膜對葡萄糖代謝的影響,發(fā)現(xiàn)各覆膜處理的SS活性均高于對照,而SPS活性均低于對照,且各處理果實中果糖和葡萄糖含量均高于對照,其中藍膜處理的葡萄糖含量最高;李夢鴿等(2016)在美人指葡萄上的研究認(rèn)為,果實生長發(fā)育過程中SS分解方向活性比較占優(yōu)勢。光質(zhì)對果實糖代謝的影響在其他果樹上也有研究,藍紅光比例1∶2對火龍果進行補光,果實中的可溶性固形物和蔗糖含量、SPS和SS活性顯著提高,紅光較少的光質(zhì)組合分解蔗糖為葡萄糖和果糖減少(陳心源等,2019);紅藍光比例6∶1的LED光源對櫻桃補光可顯著促進果實發(fā)育進程,顯著提高糖合成基因的表達和果實SS活性(李都岳和吳延軍,2023)。【本研究切入點】廣西鮮食葡萄冬果生長發(fā)育時期外界溫光變化是由強到弱的過程,果實成熟關(guān)鍵期光照條件較弱,與夏果生長發(fā)育時期的溫光變化趨勢相反(曹雄軍等,2019),光質(zhì)對廣西鮮食葡萄冬果糖代謝規(guī)律的影響鮮有報道?!緮M解決的關(guān)鍵問題】以陽光玫瑰葡萄為試驗材料,在冬果發(fā)育階段(坐果至成熟)對葡萄植株進行LED燈照射補光處理,探究廣西鮮食葡萄冬果不同生育階段的糖物質(zhì)積累代謝規(guī)律,以及光質(zhì)對果實品質(zhì)和糖物質(zhì)積累的影響,為提高葡萄冬季果實品質(zhì)提供參考依據(jù)。
1材料與方法
1.1試驗材料
試驗材料為4年生陽光玫瑰葡萄,H形架勢,株行距4 m×6 m,南北行向,連拱大棚避雨栽培,紅黃壤土質(zhì),試驗區(qū)葡萄常規(guī)管理。酶活性檢測試劑盒購自北京索萊寶科技有限公司;PrimeScriptTMRTase試劑盒購自寶日醫(yī)生物技術(shù)(北京)有限公司;Light-Cycler?480 SYBR?Green I Maste試劑盒購自瑞士Roche公司;ShimNex HE NH2柱(4.6 mm×250 mm,5μm)購自日本島津公司;果糖、蔗糖和葡萄糖標(biāo)準(zhǔn)品購自美國Sigma公司。主要儀器設(shè)備:LED紅光燈、LED藍光燈(深圳市恒春科技有限公司);普通螺旋節(jié)能白光燈(橫店集團得邦照明股份有限公司);PAL-BX/IACID2糖酸測定儀(日本ATAGO公司);PL602E電子天平(瑞士METTLER TOLEDO公司);RT-6100酶標(biāo)儀(美國Rayto公司);LightCycler?480實時熒光定量PCR儀(瑞士Roche公司);LC20AD島津液相色譜儀(日本島津公司)。
1.2試驗方法
1.2.1試驗設(shè)計試驗在位于廣西南寧市里建農(nóng)場的廣西真誠農(nóng)業(yè)有限公司葡萄基地(23°17′N,108°27′E)進行,亞熱帶季風(fēng)氣候。在陽光玫瑰葡萄冬果發(fā)育的盛花期后1周(2022年9月20日)開始補光處理,每天日出前和日落后各補光2 h,果實轉(zhuǎn)色完成后停止補光。試驗處理以每行為1個小區(qū),3次重復(fù),共設(shè)4個處理:1、LED紅光燈(RL);2、LED藍光燈(BL);3、普通螺旋節(jié)能白光燈(WL);4、不補光,作為對照(CK)。補光時,處理間用高2 m的多層黑色遮陽網(wǎng)隔開,補光用燈功率20 W,樹冠下每3 m安裝1盞補光燈,距離地面1.5 m,每次重復(fù)3株樹,每處理3次重復(fù),試驗期間田間常規(guī)管理。
1.2.2樣品采集從補光開始采樣,每周采樣1次,依照Eichhorn和Lorenz(E-L)劃分的葡萄生長階段(Keller,2015),補光期間共采樣4次:硬果期(E-L33)、果實膨大期(E-L35)、果實未完全成熟期(E-L37)、果實完全成熟期(E-L38);從每植株上隨機剪下60粒果實,果粒均來自每穗的上、中、下部位,每處理3株樹。樣品置于冰盒中帶回實驗室,每株樹的20粒果實用于測定單粒重、果??v徑、可溶性固形物、可滴定酸等指標(biāo);另40粒果實存放于-80℃冰箱,用于糖物質(zhì)含量、糖代謝相關(guān)酶活性、基因相對表達量等指標(biāo)測定。
1.2.3指標(biāo)測定利用電子天平稱量果實單粒重,游標(biāo)卡尺測量果??v徑,糖酸測定儀測定可溶性固形物和可滴定酸含量;根據(jù)酶活性檢測試劑盒的操作說明,使用酶標(biāo)儀測定SS、SPS和中性轉(zhuǎn)化酶(NI)活性;采用實時熒光定量PCR測定VvSS、VvSPS和VvNI基因表達量,用2-??Ct法(Livak and Schmittgen,2001)計算相對表達量。
使用島津液相色譜儀測定蔗糖、葡萄糖和果糖含量,參照謝林君等(2022)的方法并有所改進。液相色譜分析條件:流動相為80%乙腈水溶液,流速0.8 mL/min,柱溫25℃,進樣量20μL,檢測器為RID-20A型示差檢測器,色譜柱為ShimNex HE NH2柱(4.6 mm×250 mm,5μm)。
使用PrimeScriptTMRTase試劑盒反轉(zhuǎn)錄合成cDNA,LightCycler?480 SYBR?Green I Master試劑盒進行實時熒光定量PCR檢測,反應(yīng)體系20μL:cDNA模板1μL,上、下游引物(10μmol/L)各1μL,SYBR?GreenMix 10μL,超純水7μL。擴增程序:95℃預(yù)變性1 min;95℃10 s,55℃30 s,72℃30 s,進行40個循環(huán)。每個基因3次重復(fù)。利用Primer 6.0設(shè)計引物(表1),由奧科(武漢)生物科技有限公司合成。內(nèi)參基因為VvActin1。
1.3統(tǒng)計分析
使用Excel 2020整理試驗數(shù)據(jù),SPSS 19.0進行單因素方差分析(One-way ANOVA),以GraphPad Prism 6.0制圖。
2結(jié)果與分析
2.1補光對陽光玫瑰葡萄不同發(fā)育階段果實品質(zhì)的影響
由圖1可知,隨著陽光玫瑰葡萄發(fā)育過程的進行,果??v徑、單粒重和可溶性固形物含量逐漸提高,可滴定酸含量逐漸降低。E-L35時期果??v徑和單粒重增加較明顯,E-L37和E-L38時期果??v徑和單粒重增加幅度較小。E-L38時期白光對果粒縱徑增加的影響較大,較對照顯著提高4.94%(P<0.05,下同)。單粒重在果實成熟前各處理差異不顯著(P>0.05,下同),在E-L38時期,紅光和白光均能促進單粒重增加。果實進入成熟時期,紅光處理促進可溶性固形物含量增加,在E-L37和E-L38時期分別較對照顯著提高10.39%和11.26%。可滴定酸含量隨著果實成熟度的增加而降低,進入成熟時期,可滴定酸含量明顯減少,藍光處理對可滴定酸含量的降低有促進作用,在E-L37和E-L38時期分別較對照顯著降低15.54%和6.62%。
2.2補光對陽光玫瑰葡萄不同發(fā)育階段果實糖物質(zhì)含量的影響
由圖2可知,在E-L35時期,白光處理對陽光玫瑰葡萄果實中的果糖和葡萄糖影響較大,果實中果糖和葡萄糖含量較其他處理顯著增加。在E-L37和E-L38時期,紅光處理促進果實果糖含量增加,分別較對照顯著提高11.14%和22.93%,其他補光處理與對照差異較小。紅光處理對葡萄糖含量積累有促進作用,果實進入成熟期表現(xiàn)明顯,在E-L37和E-L38時期分別較對照顯著提高6.75%和17.51%。藍光和白光處理在果實成熟期對葡萄糖含量的積累有抑制作用,E-L38時期分別較對照顯著降低4.35%和5.62%。果實中蔗糖含量隨發(fā)育過程的進行呈先升高后降低的變化趨勢,成熟后趨于平穩(wěn),白光處理對蔗糖分解有促進作用,E-L38時期白光處理果實中蔗糖含量顯著低于對照10.12%。
2.3補光對陽光玫瑰葡萄不同發(fā)育階段果實糖代謝相關(guān)酶活性的影響
由圖3可知,在陽光玫瑰葡萄整個發(fā)育期,果實SS活性呈先升高后降低的變化趨勢。在E-L33時期,不同處理間的SS活性差異??;隨著果實發(fā)育過程的進行,紅光處理對SS活性的影響逐漸增加,在E-L35和E-L37時期,紅光處理的SS活性分別較對照顯著提高18.50%和13.39%;在E-L38時期,3個補光處理的SS活性均顯著高于對照。SPS活性在果實整個發(fā)育期均較低,呈先升高后降低的變化趨勢;在E-L38時期,紅光、藍光和白光處理均對果實SPS活性有促進作用,且紅光和藍光處理與對照存在顯著差異;在發(fā)育前期,各處理對SPS活性的影響不明顯。整個發(fā)育期,各處理對果實NI活性的影響較??;其中在E-L33、E-L35和E-L37時期,各處理的NI活性均無顯著差異;在E-L38時期,紅光處理顯著抑制NI活性增加,較對照降低17.71%。
2.4補光對陽光玫瑰葡萄不同發(fā)育階段果實糖代謝相關(guān)酶基因相對表達量的影響
由圖4可知,在陽光玫瑰葡萄果實發(fā)育初期,VvSS、VvSPS和VvNI基因相對表達量受各處理影響均不顯著。隨著果實發(fā)育過程的進行,紅光處理的VvSS基因相對表達量在E-L35、E-L37和E-L38時期分別較對照降低10.54%、86.89%和11.57%;在E-L35、E-L37、E-L38時期,藍光處理的VvSS基因相對表達量分別較對照顯著降低22.21%、47.13%和24.13%。各處理VvSPS基因相對表達量在進入果實成熟期開始表現(xiàn)不同,紅光處理的E-L37時期VvSPS基因相對表達量較對照顯著降低32.23%,藍光處理的E-L38時期VvSPS基因相對表達量較對照顯著降低22.74%。VvNI基因相對表達量在整個果實發(fā)育期呈先降低后升高的變化趨勢,紅光處理的VvNI基因相對表達量在E-L35和E-L38時期顯著低于對照,降幅分別為20.90%和28.50%;E-L38時期3個補光處理VvNI基因相對表達量均低于對照。
3討論
隨著市場需求的擴大和設(shè)施栽培的推廣,人工補光已廣泛應(yīng)用到果蔬的生長栽培中(賴瑞云等,2018;黃志午等,2022;楊肖芳等,2023;鄭曉翠等,2023;王競等,2024)。不同光質(zhì)對不同物種或同一物種不同品種植株光合作用、形態(tài)建成和生長發(fā)育的影響存在很大差異,不同光質(zhì)對植物生長、形態(tài)建成、物質(zhì)代謝及基因表達均有調(diào)控作用(時向東等,2008)。前人研究認(rèn)為,巨峰、陽光玫瑰葡萄進行補光處理后,果實的生長素、赤霉素、細胞分裂素等含量得到提高,而脫落酸含量降低,且補光對葡萄新梢生長、果實大小、產(chǎn)量、可溶性固形物含量增加及蘋果酸降解均有促進作用(白先進等,2009;時曉芳等,2021a,2021b;查倩等,2022;孔令雷等,2024)。本研究發(fā)現(xiàn),各補光處理均能促進陽光玫瑰葡萄單粒重增加,白光對果??v徑的伸長影響較大,紅光促進可溶性固形物含量增加,果實成熟期的含量顯著高于對照。藍光有助于可滴定酸含量的降低,在果實未完全成熟期差異最明顯,與張克坤等(2017)在設(shè)施栽培早熟葡萄上研究得出的結(jié)論相一致。這可能是因為不同的光環(huán)境對其形態(tài)和光合生理的影響有差異,從而導(dǎo)致光能捕捉能力和利用效率不同(Gao et al.,2013),不同光質(zhì)對植株葉片的氣孔導(dǎo)度、凈光合速率等影響有所不同,產(chǎn)生的物質(zhì)積累也存在差異(任毛飛等,2023)。
果糖、葡萄糖和蔗糖是葡萄果實中主要的可溶性糖,這些糖對果實品質(zhì)成分和風(fēng)味物質(zhì)的合成起著十分重要的作用(冷翔鵬等,2011)。光質(zhì)對高等植物的糖代謝具有調(diào)節(jié)作用(姜仲書等,2008)。本研究中,紅光促進陽光玫瑰葡萄果實果糖和葡萄糖含量的增加較明顯,與查倩等(2022)的研究結(jié)果一致。陽光玫瑰葡萄果實中蔗糖含量整體呈先升高后降低的動態(tài)變化趨勢,成熟后趨于平穩(wěn),白光處理對蔗糖分解有促進作用,說明光質(zhì)在果實發(fā)育的不同階段對蔗糖分解發(fā)揮不同作用。SS和SPS是參與蔗糖合成的關(guān)鍵酶,在果實蔗糖合成、蔗糖分解、細胞壁合成、決定果實庫強等生化過程中發(fā)揮重要作用(房經(jīng)貴和劉崇懷,2014)。補光處理能提高設(shè)施楊梅果實SPS和SS活性(任海英等,2022);LED補光處理能顯著提高甜櫻桃果實SS和SPS活性(李都岳和吳延軍,2023)。本研究中,在陽光玫瑰葡萄果實膨大期和未完全成熟期,紅光處理果實中SS活性較高;在果實完全成熟期,紅光和藍光處理均對SPS活性有促進作用,與前人研究結(jié)果(任海英等,2022;李都岳和吳延軍,2023)一致。陽光玫瑰葡萄發(fā)育過程是蔗糖逐漸被分解轉(zhuǎn)化為果糖和葡萄糖的過程,紅光處理下果實的SPS和SS活性變化規(guī)律與可溶性固形物含量積累的變化規(guī)律不一致,可能因為果實中其他有機物代謝過程與糖代謝有關(guān)聯(lián)而導(dǎo)致,有待進一步探究其關(guān)聯(lián)機理。紅藍光6∶1處理促進薄皮甜瓜CmSS01、CmSPS01基因相對表達量增加(崔曉輝,2017);補充紅光或紅藍混合光處理促進西瓜ClSPS1基因表達,提高SPS活性,達到增加西瓜蔗糖含量的目的(呂亭輝,2021)。本研究中紅光對SS和SPS活性有促進作用,但各處理對VvSS基因相對表達量有抑制作用,而對VvSPS和VvNI基因相對表達量的影響未呈現(xiàn)一定規(guī)律性。SS和SPS活性與其基因相對表達量的關(guān)系尚不明確,需從其他層面進一步解析。
4結(jié)論
紅光顯著促進陽光玫瑰葡萄冬果的可溶性固形物、果糖和葡萄糖含量積累,藍光有助于可滴定酸含量的降低。紅光、藍光和白光在果實成熟期對SS和SPS活性均有促進作用??筛鶕?jù)陽光玫瑰葡萄冬果對光照環(huán)境及果實品質(zhì)的需求選擇合適的人工光源進行補光。
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(責(zé)任編輯羅麗)