摘要:【目的】研究遮蔭及后期復(fù)光對(duì)不同類型甘薯塊根產(chǎn)量和糖類物質(zhì)的影響,以期為西南低光能輻射地區(qū)的甘薯提質(zhì)增效生產(chǎn)提供理論依據(jù)?!痉椒ā恳詢?yōu)質(zhì)食用型甘薯品種煙薯25和鮮食加工兼用型甘薯品種徐紫薯8號(hào)為試驗(yàn)材料,于大田常規(guī)密度(60000株/ha)下設(shè)3個(gè)光照/遮蔭處理:(1)全生育期自然光對(duì)照(CK);(2)栽種后50 d至收獲期持續(xù)遮光40%(S處理);(3)栽種后50 d開始40%遮蔭,持續(xù)40 d后復(fù)光(R處理),于遮蔭前1 d(T1時(shí)期)、遮蔭第40 d(T2時(shí)期)、遮蔭第80 d(T3時(shí)期)和遮蔭第120 d收獲期(T4時(shí)期)分別進(jìn)行取樣分析,對(duì)比遮蔭及復(fù)光對(duì)甘薯產(chǎn)量、干物質(zhì)積累與分配、塊根糖類物質(zhì)含量及淀粉品質(zhì)相關(guān)指標(biāo)的影響?!窘Y(jié)果】T4收獲期時(shí),煙薯25和徐紫薯8號(hào)S處理的鮮薯產(chǎn)量較CK顯著降低22.34%和10.78%(Plt;0.05,下同),R處理的鮮薯產(chǎn)量較CK降低13.28%和2.25%;S和R處理下,煙薯25單株總干重分別較CK顯著降低21.60%和18.92%,單株薯塊干重分別顯著降低和27.80%和18.77%,徐紫薯8號(hào)處理間無顯著差異(Pgt;0.05,下同)。T2時(shí)期,煙薯25和徐紫薯8號(hào)的單株葉面積和SPAD值高于CK;煙薯25比葉重顯著降低,徐紫薯8號(hào)比葉重處理間無顯著差異。T3時(shí)期,S處理下煙薯25單株葉面積和比葉重與CK無顯著差異,SPAD值顯著高于CK;徐紫薯8號(hào)S處理的單株葉面積和SPAD值顯著高于CK,比葉重顯著低于CK;R處理下煙薯25的單株葉面積和葉片SPAD值分別較CK顯著降低50.92%和5.46%,比葉重顯著增加13.03%;徐紫薯8號(hào)R處理的單株葉面積較CK顯著提高17.37%,比葉重和SPAD值與CK無顯著差異。S處理顯著提高了收獲期煙薯25的可溶性糖、還原糖和蔗糖含量,顯著降低了淀粉含量和單株淀粉產(chǎn)量;R處理的可溶性糖和淀粉含量與CK無顯著差異。徐紫薯8號(hào)收獲期S處理的可溶性糖和蔗糖含量較CK顯著降低;R處理的可溶性糖含量顯著高于S處理,但與CK無顯著差異;淀粉含量和單株淀粉產(chǎn)量在各處理間均無顯著差異。遮蔭處理使甘薯淀粉粒徑降低;T4收獲期,煙薯25淀粉溶解度和膨脹勢(shì)均表現(xiàn)為S處理gt;CKgt;R處理,且各處理間差異顯著,徐紫薯8號(hào)淀粉溶解度和膨脹勢(shì)在各處理間均無顯著差異。相關(guān)分析結(jié)果表明,煙薯25單株薯塊干重與單株總干重呈顯著或極顯著(Plt;0.01)正相關(guān),徐紫薯8號(hào)單株薯塊干重與植株物質(zhì)積累相關(guān)指標(biāo)、糖類物質(zhì)的相關(guān)系數(shù)較低。【結(jié)論】遮蔭導(dǎo)致煙薯25產(chǎn)量降低但會(huì)提升其食用品質(zhì),后期充足的光照可在保證其品質(zhì)的前提下彌補(bǔ)前期遮蔭造成的產(chǎn)量損失。徐紫薯8號(hào)對(duì)光環(huán)境具有較高的適應(yīng)性,在長
期遮蔭和光環(huán)境變化下能維持產(chǎn)量穩(wěn)定,可在光能輻射資源較低的地區(qū)種植。
關(guān)鍵詞:甘薯;遮蔭;復(fù)光;干物質(zhì)積累與分配;糖類物質(zhì);淀粉
中圖分類號(hào):S531 文獻(xiàn)標(biāo)志碼:A 文章編號(hào):2095-1191(2024)11-3242-13
Effects of shading and light restoration on yield formation andsugar substances of different types of sweet potato
LI Yong-peng1, GONG Fan-ting2, RAN Teng-fei1, XIAO Li-li1, XIA Ru1, YANG Cai1,HUANG An-zhu1, GAO Ya1, YANG Yu-quan1, TIAN Shan-jun1*
(1College of Agriculture, Guizhou University, Guiyang, Guizhou 550025, China; 2Provincial Geomatics Center ofSichuan, Ministry of Natural Resources, Chengdu, Sichuan 610041, China)
Abstract:【Objective】In order to provide theoretical basis for improving quality and efficiency of sweet potato in low light radiation area of southwestern China, the effects of shading and light restoration on root tuber yield and sugar sub‐stances of different types of sweet potato were studied.【 Method】High quality edible variety Yanshu 25 and multi-purpose variety Xuzishu 8 were used as test varieties with a planting density of 60000 plant/ha in field. Three light/shading treat‐ments were set:( 1) Natural light control during the entire planting period( CK);( 2)continuous 40% shading from 50 d post planting until harvest( S);(3) 40% shading from 50 d after planting, lasted 40 dand then restored light(R). Sam‐pling analysis was conducted 1 d before shading( T1 period), 40 d after shading( T2 period), 80 d after shading( T3 pe‐riod), and 120 d after shading( T4 period). The effects of shading and light restoration on yield, dry matter accumulation and distribution, root tuber sugar substances content and starch quality of sweet potato were compared. 【Result】During the T4 harvest period, the fresh tuber root yield of Yanshu 25 and Xuzishu 8 treated with S treatment was significantly de‐creased by 22.34% and 10.78% compared with CK( Plt;0.05, the same below), and the fresh tuber root yield of R treat‐ment was decreased by 13.28% and 2.25% compared with CK. Under S and R treatments, the total dry weight per plant of Yanshu 25 was significantly decreased by 21.60% and 18.92% compared with CK, and the root tuber dry weight per plant was significantly decreased by 27.80% and 18.77% respectively. There was no significant difference between the treat‐ments of Xuzishu 8( Pgt;0.05, the same below). At T2 period, the leaf area per plant and SPAD values of Yanshu 25 and Xuzishu 8 were higher than those of CK. The specific leaf weight of Yanshu 25 was significantly decreased, but there was no significant difference between the treatments of Xuzishu 8. At T3 period, the leaf area and specific leaf weight of Yanshu 25 were not significantly different from CK under S treatment, and the SPAD value was significantly higher than CK. The leaf area per plant and SPAD value of Xuzishu 8 treated with S treatment were significantly higher than CK, and the specific leaf weight was significantly lower than CK. Under R treatment, the leaf area per plant and leaf SPAD value of Yanshu 25 were significantly decreased by 50.92% and 5.46% compared with CK respectively, and specific leaf weight was significantly increased by 13.03%. Compared with CK, the leaf area per plant of Xuzishu 8 treated with R treatment was significantly increased by 17.37%, and the specific leaf weight and SPAD value were not significantly different from CK. S treatment significantly increased the contents of soluble sugar, reducing sugar and sucrose in Yanshu 25, and sig‐nificantly decreased the starch content and starch yield per plant. The contents of soluble sugar and starch in R treatment were not significantly different from those in CK. The contents of soluble sugar and sucrose in S treatment of Xuzishu 8 during harvesting period was significantly decreased than that in CK. The soluble sugar content of R treatment was signifi‐cantly higher than that of S treatment, but there was no significant difference between R treatment and CK. There was no significant difference in starch content and starch yield per plant among different treatments. Shading treatment reduced the particle size of sweet potato starch. At T4 harvest period, the solubility and swelling power of starch of Yanshu 25 were as follows: S treatment gt; CK gt; R treatment, and there were significant differences among all treatments. There were no significant differences in solubility and swelling power of starch of Xuzishu 8 among all treatments. The results of cor‐relation analysis showed that the root tuber dry weight per plant of Yanshu 25 was significantly or extremely significantly (Plt;0.01) positively correlated with the total dry weight per plant, and the correlation coefficients between root tuber dry weight per plant and plant material accumulation, sugar substance of Xuzizhu 8 was low. 【Conclusion】Shading reduces the yield of Yanshu 25 but improves edible quality. Sufficient light in the later period can make up for the yield loss caused by the early shading under the premise of ensuring its quality. Xuzishu 8 has high adaptability to the light environment, and can maintain stable yield under long-term shading and light environment changes. It can be planted in areas with low
light energy radiation resources.
Key words: sweet potato; shading; light restoration; dry matter accumulation and distribution; sugar substances;
starch
Foundation items: Guizhou Science and Technology Support Plan Projec(t QKHZC〔2022〕Zhongdian 013,QKHZC〔2022〕Zhongdian 019-05)
0 引言
【研究意義】甘薯是重要的糧飼經(jīng)兼用作物,在保障我國糧食安全中大有可為(李強(qiáng)等,2022),西南地區(qū)是我國甘薯的重要產(chǎn)地之一。但甘薯是喜光作物,西南產(chǎn)區(qū)中的貴州省是太陽輻射低值區(qū)域,當(dāng)?shù)靥栞椛淞亢腿照諘r(shí)數(shù)均處于全國較低水平且時(shí)空分布不均(王凌鈺,2023),嚴(yán)重制約了甘薯的高產(chǎn)優(yōu)質(zhì)生產(chǎn)。隨著鄉(xiāng)村振興戰(zhàn)略的推進(jìn),市場對(duì)甘薯的高值化利用提出了更高要求,甘薯的用途不斷向多樣化和專用化發(fā)展(馬代夫等,2021),薯干率及可溶性糖和淀粉含量等理化性質(zhì)成為優(yōu)質(zhì)甘薯生產(chǎn)的重要評(píng)價(jià)指標(biāo)。因此,基于甘薯生產(chǎn)中遭受的光能輻射不足及時(shí)空分布不均對(duì)甘薯產(chǎn)量和品質(zhì)的影響,研究遮蔭及復(fù)光對(duì)不同類型甘薯產(chǎn)量、糖類物質(zhì)和淀粉品質(zhì)的影響,可為低光能輻射條件下優(yōu)質(zhì)甘薯的生產(chǎn)提供理論支撐,助力產(chǎn)業(yè)鏈條拓展。【前人研究進(jìn)展】關(guān)于甘薯光蔭蔽的研究早期主要集中在形態(tài)發(fā)育、光合生理和產(chǎn)量等方面。遮蔭造成的甘薯塊根減產(chǎn)是由植物生物量積累減少和光合產(chǎn)物分配改變共同引起的,生長中后期光照不足會(huì)造成更大的產(chǎn)量損失(Oswald et al.,1995)。遮蔭下,甘薯主蔓長、地上重量和根冠比顯著增高,塊根產(chǎn)量顯著下降(王慶美,2008)。侯夫云等(2013)研究表明,短期遮蔭下甘薯葉片的超氧化物歧化酶(SOD)和過氧化氫酶(CAT)活性下降,過氧化物酶(POD)活性和丙二醛(MAD)含量升高,但長期遮蔭下甘薯葉片抗氧化防御系統(tǒng)相關(guān)指標(biāo)變化存在明顯的品種間差異。李韋柳等(2017)以桂經(jīng)薯2號(hào)為材料,研究不同遮陰處理對(duì)甘薯生長的影響,結(jié)果表明,遮陰導(dǎo)致甘薯葉片增大,葉綠素含量升高,鮮薯和淀粉產(chǎn)量及葉綠素a/b降低;且遮陰對(duì)甘薯的影響隨遮陰程度的增加而加劇。此外,也有學(xué)者基于農(nóng)藝性狀、產(chǎn)量、光合生理指標(biāo)等對(duì)不同品種甘薯進(jìn)行耐蔭性評(píng)價(jià)和耐蔭性品種篩選。蔣亞(2020)對(duì)29個(gè)甘薯品種(系)進(jìn)行耐蔭性評(píng)價(jià),結(jié)果表明強(qiáng)耐蔭性品種植株的形態(tài)發(fā)育、光合生理和抗逆性指標(biāo)受蔭蔽的影響較小,且復(fù)光后莖蔓生長的恢復(fù)和光合能力提高等更具優(yōu)勢(shì)。近年來,轉(zhuǎn)錄組和代謝組等技術(shù)被應(yīng)用到甘薯弱光響應(yīng)機(jī)制研究中。楊津(2022)對(duì)弱光脅迫下2個(gè)不同耐蔭性甘薯品種進(jìn)行轉(zhuǎn)錄組分析,發(fā)現(xiàn)在遮蔭環(huán)境下,葉綠體膜醌氧化還原酶同系物相關(guān)基因和黃酮醇合酶/黃烷酮3-羥化酶相關(guān)基因表達(dá)量在2個(gè)甘薯品種葉片中均顯著下調(diào);涉及光合作用中鐵氧還蛋白相關(guān)基因在耐弱光脅迫型品種中顯著下調(diào),在弱光脅迫敏感型品種中顯著上調(diào)。王雁楠等(2023)結(jié)合轉(zhuǎn)錄組和代謝組解析遮陰脅迫下甘薯代謝響應(yīng)途徑,發(fā)現(xiàn)苯丙素合成途徑、糖代謝相關(guān)途徑、鞘脂和精氨酸代謝途徑是甘薯的主要弱光響應(yīng)代謝途徑。甘薯品質(zhì)研究方面,王慶美(2008)研究表明,不同遮蔭程度均會(huì)導(dǎo)致甘薯薯干率和總淀粉含量下降、可溶性糖含量增高,遮蔭對(duì)甘薯直/支鏈淀粉、可溶性蛋白和花青素含量的影響存在品種間差異;王藝丹(2022)研究表明,20%遮光可在產(chǎn)量不降低的前提下提高紫薯塊根中可溶性糖、花色苷和類黃酮含量?!颈狙芯壳腥朦c(diǎn)】當(dāng)前對(duì)弱光脅迫下甘薯形態(tài)發(fā)育、產(chǎn)量形成和生理響應(yīng)已有較深入研究,但品質(zhì)相關(guān)研究較少且主要集中在紫肉甘薯品種。當(dāng)前市場甘薯用途向多樣化和專用化發(fā)展,作為淀粉及其衍生產(chǎn)品加工原料的甘薯需較高的淀粉含量和薯塊干物率,鮮食和食品加工型甘薯要求具有較高的糖含量和相對(duì)更低的干物率,因此在產(chǎn)量研究的基礎(chǔ)上探究遮蔭及復(fù)光對(duì)不同類型甘薯品質(zhì)的影響,對(duì)弱光環(huán)境下優(yōu)質(zhì)甘薯的生產(chǎn)具有切實(shí)的參考價(jià)值?!緮M解決的關(guān)鍵問題】針對(duì)貴州省甘薯生產(chǎn)過程中弱光環(huán)境造成的產(chǎn)量降低和品質(zhì)變化,以煙薯25和徐紫薯8號(hào)為試驗(yàn)材料,采用大田栽培試驗(yàn),設(shè)人工遮蔭和復(fù)光處理,研究遮蔭及后期復(fù)光對(duì)甘薯塊根產(chǎn)量積累、糖類物質(zhì)和淀粉性質(zhì)的影響,以期為西南低光能輻射區(qū)域的甘薯提質(zhì)增效生產(chǎn)提供理論依據(jù)。
1 材料與方法
1. 1 試驗(yàn)地概況及試驗(yàn)品種
試驗(yàn)于2022年5—11月在貴州省思南縣雙塘街道何家寨村(27°59′3.58″N,108°12′30.96″E)開展。試驗(yàn)地年均氣溫17.4 ℃,年日照時(shí)數(shù)1291.5 h;前茬作物為油菜;土壤類型為黃壤,pH 6.95,有機(jī)質(zhì)含量22.83 g/kg,堿解氮含量126.3 mg/kg,有效磷含量18.83 mg/kg,速效鉀含量85.07 mg/kg。
供試甘薯品種為煙薯25和徐紫薯8號(hào)。煙薯25為優(yōu)質(zhì)食用型甘薯品種,徐紫薯8號(hào)為鮮食加工兼用型紫肉甘薯品種,種苗由國家甘薯種質(zhì)資源試管苗庫(徐州)提供。
1. 2 試驗(yàn)方法
田間試驗(yàn)采用二因素裂區(qū)試驗(yàn)設(shè)計(jì),主區(qū)為光照/遮蔭處理,副區(qū)為品種。主區(qū)設(shè)3個(gè)光照/遮蔭處理:(1)全生育期自然光對(duì)照(CK);(2)栽種后50 d至收獲期持續(xù)遮光40%(S處理);(3)栽種后50 d開始40%遮蔭,持續(xù)40 d后復(fù)光(R處理)。肥料用量為N 110 kg/ha、P2O5 40 kg/ha、K2O 120 kg/ha,全部作基肥施用,以復(fù)合肥(15-5-20)、尿素和過磷酸鈣混勻,均勻撒施后深翻入地并起壟。小區(qū)長4 m、寬2 m,壟距0.8 m、壟高約25 cm,密度60000株/ha。選取長勢(shì)一致的薯苗斜插種植,入土3節(jié)。采用黑色遮陽網(wǎng)進(jìn)行遮蔭,遮陽網(wǎng)距地面高度1.5 m,遮蔭小區(qū)四周同樣遮擋遮陽網(wǎng)。利用照度計(jì)比對(duì)遮陽網(wǎng)內(nèi)外光照,通過調(diào)整遮陽網(wǎng)綁扎松緊程度,使網(wǎng)內(nèi)各區(qū)域遮蔭程度保持一致。其他管理措施同當(dāng)?shù)馗弋a(chǎn)田生產(chǎn)。1. 3 測(cè)定項(xiàng)目及方法
1. 3. 1 樣品采集與處理 于遮蔭前1 d(T1時(shí)期)、遮蔭第40 d(T2時(shí)期)、遮蔭第80 d(T3時(shí)期)和遮蔭第120 d收獲期(T4時(shí)期)分別進(jìn)行取樣,每小區(qū)取樣6株。其中3株按器官分開,進(jìn)行干物質(zhì)重分析。剩余3株塊根清洗后,液氮速凍,-80 ℃保存,用于糖類指標(biāo)測(cè)定。
1. 3. 2 塊根鮮重、植株干重及薯塊干物率測(cè)定 鮮樣先經(jīng)105 ℃殺青15 min,再于80 ℃烘干至恒重,稱取各器官(根、塊根、莖、葉、葉柄)。干重并計(jì)算植株的單株總干重和地上部干重(莖、葉、葉柄)。烘干前稱取塊根鮮重,計(jì)算薯塊干物率,薯塊干物率(%)=烘干后塊根干重/烘干前塊根鮮重×100。
1. 3. 3 葉片SPAD值、葉面積和比葉重測(cè)定 使用SPAD-502葉綠素含量測(cè)定儀,田間測(cè)定T1、T2、T3時(shí)期植株主蔓第 4片成熟葉中部距葉脈 2 cm處SPAD值,每小區(qū)測(cè)定3株。使用Li-3100C掃描葉面積儀測(cè)定甘薯單株全部綠葉葉面積,計(jì)算比葉重,比葉重(g/m2)=單株綠葉干重/單株綠葉面積。1. 3. 4 糖類物質(zhì)測(cè)定 糖類物質(zhì)含量測(cè)定參考王三根(2017)的方法。取樣品1 g,加入5 mL 80%乙醇充分研磨后全部移入15 mL離心管,80 ℃水浴提取30 min,離心,上清液倒入20 mL離心管,沉淀復(fù)提2次,合并上清液并定容,為待測(cè)總可溶性糖提取液,沉淀烘干備用??偪扇苄蕴翘崛∫合♂屩吝m宜濃度后,采用蒽酮—硫酸法測(cè)定可溶性糖含量,DNS法測(cè)定還原糖含量,間苯二酚法測(cè)定蔗糖含量。烘干后的沉淀以鹽酸水解—DNS法測(cè)定淀粉含量。糖類物質(zhì)均為鮮基含量。1. 3. 5 淀粉粒徑分布及形貌特征分析 參考Abe‐gunde等(2013)的方法提取淀粉,粒徑測(cè)定采用Bet‐tersize 2000E激光粒度儀,以純水為分散介質(zhì),超聲分散5 min后測(cè)定,重復(fù)3次,以平均值為最終測(cè)定結(jié)果。樣品粘到導(dǎo)電膠上,噴金30 s后使用TESCAN MIRA LMS掃描電子顯微鏡拍攝樣品形貌。1. 3. 6 淀粉溶解度和膨脹勢(shì) 參考Chio等(2009)的方法,取淀粉樣品0.5 g于50 mL離心管,加入30 mL蒸餾水,90 ℃水浴30 min;冷卻至室溫后1500 g離心30 min,上清液倒入已知質(zhì)量的鋁盒,并稱取沉淀濕重。上清液在105 ℃下干燥16 h后稱取上清液干重。溶解度和膨脹勢(shì)計(jì)算公式如下:溶解度(%)=上清液干重/樣品重量×100膨脹勢(shì)(g/g)=沉淀濕重×100[/樣品重量×(100-"""""""""""""""""""""" 溶解度)]
1. 4 統(tǒng)計(jì)分析采用Excel 2010進(jìn)行數(shù)據(jù)整理和統(tǒng)計(jì)繪圖,利用IBM SPSS Statistics 25.0進(jìn)行多重比較,以O(shè)rigin 2019繪制淀粉粒徑分布圖,采用R 4.1.3進(jìn)行相關(guān)分析和繪圖。
2 結(jié)果與分析
2. 1 遮蔭及復(fù)光對(duì)鮮薯產(chǎn)量的影響由表1可知,T2時(shí)期,田間遮蔭40 d導(dǎo)致甘薯鮮薯產(chǎn)量明顯降低;T3時(shí)期,S處理煙薯25鮮薯產(chǎn)量較CK顯著降低19.99%(Plt;0.05,下同),徐紫薯8號(hào)鮮薯產(chǎn)量與CK無顯著差異(Pgt;0.05,下同);T4收獲期,S處理煙薯25和徐紫薯8號(hào)鮮薯產(chǎn)量分別較CK顯著降低22.34%和10.78%。遮蔭后復(fù)光導(dǎo)致短期內(nèi)鮮薯產(chǎn)量較S處理進(jìn)一步降低,T3時(shí)期R處理煙薯25和徐紫薯8號(hào)鮮薯產(chǎn)量均低于CK和S處理;但隨著復(fù)光時(shí)間延長,R處理下鮮薯產(chǎn)量逐漸增長,到T4收獲期,R處理煙薯25和徐紫薯8號(hào)鮮薯產(chǎn)量分別較CK降低13.28%和2.25%,但均高于S處理,最終鮮薯產(chǎn)量表現(xiàn)為CKgt;R處理gt;S處理。從薯塊干物率來看,T2時(shí)期,各處理的薯塊干物率差異不顯著;T3時(shí)期,隨著鮮薯產(chǎn)量的增加,薯塊干物率較T2時(shí)期下降,該時(shí)期煙薯25各處理的薯塊干物率無顯著差異,但S處理和CK均低于R處理,徐紫薯8號(hào)S處理與CK的薯塊干物率無顯著差異,但二者均顯著低于R處理;T4時(shí)期,2個(gè)甘薯品種的薯塊干物率較T3時(shí)期提高,煙薯 25表現(xiàn)為 S和 R處理顯著低于CK,徐紫薯8號(hào)表現(xiàn)為S處理gt;R處理gt;CK,S處理與CK間差異顯著。
2. 2 遮蔭及復(fù)光對(duì)甘薯葉片性狀的影響
由圖1可知,遮蔭40 d后,煙薯25單株葉面積和葉片SPAD值均較CK增大,而比葉重較CK顯著降低。同時(shí)期,徐紫薯8號(hào)的單株葉面積、比葉重和SPAD值均高于CK,其中SPAD值與CK差異顯著。T2到T3時(shí)期,2個(gè)甘薯品種的單株葉面積大幅降低,T3時(shí)期,煙薯25 S處理的單株葉面積、比葉重與CK差異不顯著,SPAD值顯著提高,徐紫薯8號(hào)S處理的單株葉面積和SPAD值分別較CK顯著增加32.36%和9.08%,比葉重較CK顯著降低27.13%。遮蔭后復(fù)光導(dǎo)致T3時(shí)期煙薯25單株葉面積分別較CK和S處理顯著降低50.92%和45.29%,SPAD值分別較CK和S處理顯著降低5.46%和9.54%,且葉面積較T2時(shí)期降幅明顯高于CK和S處理,比葉重分別較CK和S處理顯著增加13.03%和11.14%;T3時(shí)期徐紫薯8號(hào)R處理單株葉面積較CK顯著提高17.37%,但顯著低于S處理,比葉重和SPAD值均與CK無顯著差異,但比葉重顯著高于S處理,SPAD值顯著低于S處理。
2. 3 遮蔭及復(fù)光對(duì)甘薯干物質(zhì)積累和分配的影響
由圖2可知,遮蔭和復(fù)光會(huì)導(dǎo)致甘薯干物質(zhì)分配在處理間表現(xiàn)出明顯差異。遮蔭40 d后,煙薯25單株總干重較CK降低5.45%~8.76%,單株薯塊干重較CK顯著降低13.14%~15.06%,在干物質(zhì)分配比例上表現(xiàn)為遮蔭處理下塊根分配比例減小,莖分配比例增大。T3到T4時(shí)期,CK和S處理單株總干重略有下降但單株薯塊干重仍然提高。T4時(shí)期,S處理煙薯25單株總干重和單株薯塊干重分別較CK顯著降低21.60%和27.80%。在干物質(zhì)分配比例上,S處理塊根占比在T3和T4時(shí)期分別為55.98%和64.97%,CK分別為66.51%和72.21%。表明遮蔭對(duì)煙薯25產(chǎn)量的負(fù)面影響至少來自于物質(zhì)積累效率下降和分配失衡兩方面。復(fù)光后,T3時(shí)期R處理煙薯25單株總干重顯著低于CK和S處理,甚至低于T2時(shí)期,但其塊根干物質(zhì)占比較高,單株薯塊干重與S處理無顯著差異。T4時(shí)期,R處理下單株總干重和塊根占比均較T3時(shí)期提高,單株總干重和單株薯塊干重分別較CK顯著降低18.92%和18.77%,單株薯塊干重顯著高于S處理。
遮蔭和復(fù)光對(duì)徐紫薯8號(hào)單株總干重和塊根物質(zhì)積累的影響與煙薯25存在差異。徐紫薯8號(hào)各處理下單株總干重和單株薯塊干重始終呈上升趨勢(shì),且各時(shí)期單株總干重處理間均無顯著差異,但器官物質(zhì)分配比例在各處理間存在明顯差異。遮蔭40 d后,S和R處理塊根占比低于CK,與煙薯25相似。
到T3時(shí)期,S處理塊根物質(zhì)占比和單株薯塊干重均顯著高于CK和R處理。到T4收獲期,S處理塊根干物質(zhì)占比低于CK和R處理,處理間單株薯塊干重?zé)o顯著差異??梢?,復(fù)光后徐紫薯8號(hào)各器官干物質(zhì)分配特征和單株薯塊干重與S處理出現(xiàn)明顯區(qū)別,與CK差異逐漸降低。
2. 4 遮蔭及復(fù)光對(duì)甘薯塊根糖類物質(zhì)的影響
由圖3可知,遮蔭導(dǎo)致T2和T3時(shí)期甘薯塊根可溶性糖含量較CK提高,但T4時(shí)期,煙薯25的S處理顯著高于CK,徐紫薯8號(hào)的S處理顯著低于CK。復(fù)光后,煙薯25在R處理下可溶性糖含量與CK基本持平,徐紫薯8號(hào)在T3時(shí)期R處理下的可溶性糖含量顯著高于CK和S處理,T4時(shí)期降低至與CK無顯著差異。S處理下T2和T4時(shí)期煙薯25的還原糖含量顯著高于CK,T3時(shí)期各處理間無顯著差異,徐紫薯8號(hào)各時(shí)期S處理的還原糖含量均顯著高于CK。復(fù)光后,T3時(shí)期煙薯25和徐紫薯8號(hào)R處理還原糖含量與CK無顯著差異,T4時(shí)期則顯著高于CK,且徐紫薯8號(hào)R處理還原糖含量顯著高于S處理。遮蔭導(dǎo)致2個(gè)品種T2時(shí)期甘薯塊根蔗糖含量較CK顯著降低,到T3時(shí)期S處理與CK的蔗糖含量無顯著差異,T4時(shí)期S處理下煙薯25的蔗糖含量顯著高于CK,徐紫薯8號(hào)則顯著低于CK。R處理下,T3和T4時(shí)期煙薯25的蔗糖含量均低于CK和S處理,徐紫薯8號(hào)在T3時(shí)期顯著高于CK和S處理,T4時(shí)期顯著高于S處理,與CK無顯著差異??傮w而言,遮蔭導(dǎo)致收獲期甘薯還原糖含量較CK升高,可溶性糖和蔗糖含量變化則因品種而異,復(fù)光可導(dǎo)致收獲期塊根可溶性總糖含量趨近于CK,但還原糖含量較S處理進(jìn)一步提升。
2. 5 遮蔭及復(fù)光對(duì)甘薯淀粉積累及性質(zhì)的影響
2. 5. 1 遮蔭及復(fù)光處理下甘薯塊根淀粉積累 由圖4可知,遮蔭40 d后,煙薯25塊根淀粉含量顯著高于CK,徐紫薯8號(hào)塊根淀粉含量顯著低于CK;T3時(shí)期,2個(gè)品種S處理的淀粉含量與CK無顯著差異且均低于T2時(shí)期;T4收獲期,煙薯25 S處理的淀粉含量較CK顯著降低16.29%,徐紫薯8號(hào)S處理與CK無顯著差異。遮蔭后復(fù)光導(dǎo)致2個(gè)品種T3時(shí)期淀粉含量顯著高于S處理和CK,隨后含量降低,到T4時(shí)期降至與CK無顯著差異。
單株淀粉產(chǎn)量方面,T2時(shí)期,煙薯25單株淀粉產(chǎn)量在各處理間無顯著差異,但遮蔭導(dǎo)致徐紫薯8號(hào)單株淀粉產(chǎn)量顯著降低。隨著遮蔭的持續(xù)進(jìn)行和復(fù)光的推進(jìn),到T3和T4時(shí)期,S和R處理下煙薯25的單株淀粉產(chǎn)量低于CK。T3時(shí)期,徐紫薯8號(hào)S處理的單株淀粉產(chǎn)量與CK無顯著差異,R處理則顯著低于CK和S處理,T4時(shí)期CK略高于S和R處理,但差異不顯著。
2. 5. 2 遮蔭及復(fù)光后收獲期甘薯淀粉粒徑分布和形貌特征 由表2可知,淀粉粒徑存在明顯的品種間差異。煙薯25 CK的塊根淀粉粒徑D10和D90分別為1.82和31.08 μm,R處理塊根淀粉粒徑D10和D90分別為1.77和31.62 μm,即占據(jù)80%體積的淀粉粒粒徑分布集中在1.77~31.62 μm,相較于CK的粒度分布更加分散,S處理下該范圍在1.85~28.87 μm,相對(duì)于CK粒徑范圍更加集中。徐紫薯8號(hào)CK 80%體積的淀粉粒粒徑分布集中在1.95~26.57 μm,變化幅度小于煙薯25的CK;S處理下80%體積的淀粉粒粒徑分布集中在1.99~26.46 μm,較CK更集中;而R處理下80%體積的淀粉粒粒徑分布集中在2.01~25.63 μm,分布變化范圍較CK和R處理更小。
圖5為煙薯25和徐薯紫8號(hào)淀粉體積粒徑分布頻率圖。煙薯25各處理下粒徑分布峰值分別在1.47和17.58 μm處,較多的粒徑分布在第2個(gè)峰值附近的11.2~27.6 μm范圍內(nèi),CK、S和R處理下該范圍內(nèi)淀粉粒徑分布累積頻率分別為47.18%、49.78%和47.32%。徐紫薯8號(hào)CK和R處理下粒徑分布峰值與煙薯25相同,S處理下第2個(gè)峰值在14.03 μm處;粒徑分布頻率主要集中在8.94~22.03 μm范圍內(nèi),該范圍內(nèi)CK、S和R處理的分布頻率分別為52.68%、51.27%和54.59%。
從圖6掃描電子顯微鏡拍攝的照片可觀察到不同處理下甘薯塊根淀粉形貌特征。甘薯淀粉形態(tài)上總體表現(xiàn)為球體或半球體,部分顆粒具有1個(gè)或多個(gè)凹陷。煙薯25大顆粒淀粉(≥10 μm)呈具多個(gè)凹陷的半球體狀或小半個(gè)球體狀,其凹陷多處于半球底面?zhèn)确剑姑鏋楣饣∶?,凹面有不?guī)則起伏。小顆粒淀粉()中除了具有與大顆粒類似形貌,部分顆粒呈光滑球體和底部具單一凹陷的半球體。CK和R處理比S處理有更多具棱不規(guī)則顆粒。徐紫薯8號(hào)與煙薯25相比,底部具單一凹陷的半球體顆粒更多,尤其是CK和R處理。S處理與CK和R處理相比,小顆粒淀粉所占比例更大,且多呈底部側(cè)面有多個(gè)凹陷的半球體狀。
2. 5. 3 遮蔭及復(fù)光處理后甘薯淀粉溶解度及膨脹勢(shì) 由圖7可看出,T2時(shí)期,煙薯25和徐紫薯8號(hào)塊根淀粉溶解度均較CK顯著降低;T2到T4時(shí)期,煙薯25的CK和R處理下淀粉溶解度先增高后降低,S處理下表現(xiàn)為T2到T3時(shí)期大幅增高,T3到T4期間基本維持穩(wěn)定;T3時(shí)期,煙薯25 CK和S處理的淀粉溶解度顯著低于R處理,T4時(shí)期S處理的淀粉溶解度顯著高于CK和R處理。徐紫薯8號(hào)淀粉溶解度在13.8%~18.0%,CK的淀粉溶解度在各時(shí)期均高于S和R處理,且S和R處理間無顯著差異,且各時(shí)期間變化幅度明顯小于煙薯25。
由圖7還可看出,2個(gè)品種T2時(shí)期的淀粉膨脹勢(shì)在各處理間均無顯著差異,T3時(shí)期表現(xiàn)為CKlt;S處理lt;R處理;到T4時(shí)期,煙薯25淀粉膨脹勢(shì)表現(xiàn)為S處理gt;CKgt;R處理,處理間差異顯著,徐紫薯8號(hào)處理間無顯著差異??傮w而言,遮蔭和復(fù)光會(huì)導(dǎo)致煙薯25淀粉溶解度和膨脹勢(shì)表現(xiàn)出較大變化,徐紫薯8號(hào)則更穩(wěn)定。
2. 6 甘薯產(chǎn)量指標(biāo)、干物質(zhì)分配及糖類物質(zhì)的相關(guān)性
圖8為甘薯產(chǎn)量指標(biāo)、干物質(zhì)分配及塊根糖類物質(zhì)的相關(guān)性熱圖。圖8-A顯示,在單株生物量積累上,煙薯25各時(shí)期單株總干重和單株薯塊干重相互之間,以及其各自與T3和T4時(shí)期的根、葉、葉柄、地上部干重和各時(shí)期單株薯塊干重呈正相關(guān),尤其與各時(shí)期單株薯塊干重幾乎均呈顯著或極顯著(Plt;0.01,下同)相關(guān),即單株薯塊干重和植株物質(zhì)積累間表現(xiàn)為同步增長。糖類物質(zhì)方面,T2和T3時(shí)期單株總干重、單株薯塊干重與同時(shí)期蔗糖含量呈正相關(guān),與還原糖、可溶性糖和淀粉含量則呈負(fù)相關(guān)。到T4時(shí)期則發(fā)生改變,該時(shí)期單株總干重、單株薯塊干重與同時(shí)期蔗糖、還原糖和可溶性糖含量呈負(fù)相關(guān),與淀粉含量呈正相關(guān)。淀粉積累方面,單株淀粉產(chǎn)量與單株總干重和單株薯塊干重呈正相關(guān)。圖8-B顯示,對(duì)于徐紫薯8號(hào),同時(shí)期下,T2時(shí)期單株總干重與薯塊干物率、地上部干重、莖干重均呈顯著正相關(guān),與鮮薯產(chǎn)量呈顯著負(fù)相關(guān)。鮮薯產(chǎn)量與地上部、莖、葉干重呈顯著或極顯著負(fù)相關(guān),即同樣或更高的總物質(zhì)量積累下,地上部的旺盛生長可能導(dǎo)致更低的鮮薯產(chǎn)量。T3時(shí)期鮮薯產(chǎn)量則與薯塊干物率、蔗糖含量、淀粉含量、莖干重呈顯著或極顯著負(fù)相關(guān)。
3 討論
3. 1 遮蔭及復(fù)光對(duì)不同類型甘薯產(chǎn)量的影響
高光合產(chǎn)物積累量和合理的物質(zhì)分配比例有利于保障作物產(chǎn)量。遮蔭弱光條件下作物群體光能輻射截獲量減少,光合作用受到光環(huán)境因素的制約,光合產(chǎn)物積累受限(劉鑫等,2023)。同時(shí),在對(duì)弱光環(huán)境的適應(yīng)性改變中,植物的形態(tài)建成和物質(zhì)分配規(guī)律被打破(劉典三,2014;侯月爽等,2023)。針對(duì)甘薯生產(chǎn)中面臨的弱光逆境,前人使用不同指標(biāo)進(jìn)行甘薯耐蔭性評(píng)價(jià),探究甘薯耐蔭性評(píng)價(jià)指標(biāo)的同時(shí),證明了甘薯弱光響應(yīng)的品種間差異(呂長文等,2023)。本研究中,遮蔭后煙薯25單株總干重、鮮薯產(chǎn)量和單株薯塊干重顯著低于自然光對(duì)照,塊根產(chǎn)量降低幅度大于總生物量降低幅度,說明遮蔭對(duì)煙薯25產(chǎn)量的影響是植株物質(zhì)積累總量減少和塊根分配比例減少共同作用的結(jié)果;相同處理下,徐紫薯8號(hào)和煙薯25的不同產(chǎn)量變化表明甘薯在遮蔭和復(fù)光下存在品種間差異。吳雨珊(2021)研究發(fā)現(xiàn)大豆在蔭蔽后復(fù)光存在系統(tǒng)性補(bǔ)償,以彌補(bǔ)蔭蔽造成的生長損失。本研究發(fā)現(xiàn),復(fù)光后,短期內(nèi)甘薯生長和產(chǎn)量積累相較于持續(xù)遮蔭和全生育期自然光對(duì)照均表現(xiàn)出劣勢(shì);隨著復(fù)光時(shí)間延長,到收獲期,復(fù)光處理下甘薯產(chǎn)量表現(xiàn)出高于持續(xù)遮蔭但低于自然光對(duì)照的結(jié)果,后期產(chǎn)量增長率高于另外2個(gè)處理,說明經(jīng)歷弱光環(huán)境的甘薯在恢復(fù)并適應(yīng)自然光環(huán)境后在產(chǎn)量上同樣存在補(bǔ)償效應(yīng)。此外,同條件下徐紫薯8號(hào)產(chǎn)量變化幅度小,即以產(chǎn)量而言,徐紫薯8號(hào)對(duì)光環(huán)境變化具有更高的適應(yīng)性。
3. 2 遮蔭及復(fù)光對(duì)不同類型甘薯光合能力的影響
葉片是植物最主要的光合器官。李彩斌和郭華春(2017)的馬鈴薯遮蔭試驗(yàn)、雷怡等(2023)的大豆弱光脅迫試驗(yàn)中存在遮蔭后葉面積指數(shù)增高的品種。高等植物為適應(yīng)弱光逆境,表現(xiàn)出增加葉面積以提升光能截獲能力的響應(yīng)機(jī)制。本研究也表明,在甘薯形態(tài)建成旺盛的生長中期,遮蔭處理下甘薯功能葉的葉面積高于自然光對(duì)照,遮蔭導(dǎo)致功能葉SPAD值提高,與蔣亞(2020)的研究中遮蔭處理下甘薯葉片葉綠素含量升高的結(jié)果相同。而王慶美(2008)的研究則表明遮蔭對(duì)甘薯葉綠素含量的影響因品種和遮蔭程度會(huì)表現(xiàn)出高于或低于自然光對(duì)照。遮蔭下甘薯葉片柵欄組織和海綿組織厚度下降,葉片厚度減小(楊津,2022),導(dǎo)致單位葉面積下葉鮮重降低。由此可見,在通過葉綠素含量討論遮蔭下植物光合能力時(shí),還需綜合考慮單位葉面積和單位質(zhì)量下的葉綠素含量。Wang等(2014)的研究結(jié)果表明,與自然光照下形成的葉片相比,遮蔭下形成的甘薯葉片單位葉面積上的光合效率更低,表明比葉重與葉片光合能力具有密切關(guān)聯(lián)。本研究中,遮蔭40 d后煙薯25比葉重顯著低于自然光對(duì)照,是間接保障植株產(chǎn)量的前提。
3. 3 遮蔭及復(fù)光對(duì)不同類型甘薯品質(zhì)的影響
本研究中,煙薯25在收獲期具有較低的薯塊干物率和較高的可溶性糖含量,與史春余等(2008)、Xu等(2023)對(duì)優(yōu)質(zhì)鮮食型甘薯的評(píng)價(jià)相符;而作為食用加工兼用型的徐紫薯8號(hào)則具有更高的薯塊干物率和較低的可溶性糖含量。遮蔭導(dǎo)致煙薯25鮮薯產(chǎn)量和薯塊干物率降低,可溶性糖含量增高,食用品質(zhì)提升。復(fù)光后,短期內(nèi)產(chǎn)量降低,到收獲時(shí),R處理下可溶性糖含量不低于自然光對(duì)照,鮮薯產(chǎn)量較S處理增高。由此可知,生長后期充足的光照可在保障煙薯25品質(zhì)的前提下,彌補(bǔ)前期光能不足造成的產(chǎn)量損失。淀粉的形貌和粒徑與部分加工品質(zhì)具有顯著相關(guān)性(張令文等,2021),進(jìn)而影響淀粉加工產(chǎn)品的品質(zhì)。Chen等(2003)研究認(rèn)為小顆粒淀粉()制作的粉條品質(zhì)優(yōu)于以大顆粒淀粉或不同粒徑顆?;旌系矸蹫樵现谱鞯姆蹢l。余樹璽等(2015)研究表明,甘薯淀粉粒徑、溶解度等指標(biāo)與粉條品質(zhì)呈負(fù)相關(guān),膨脹勢(shì)等與粉條品質(zhì)呈正相關(guān)。本研究中,遮蔭導(dǎo)致徐紫薯8號(hào)淀粉粒徑降低,且生長后期復(fù)光進(jìn)一步降低淀粉粒徑。同時(shí)遮蔭處理下,徐紫薯8號(hào)淀粉溶解度降低,膨脹勢(shì)增高。復(fù)光后,在淀粉溶解度與遮蔭無差異的背景下,膨脹勢(shì)進(jìn)一步明顯升高。由此可知,夏季遮蔭及生長后期保障充足的光照,有利于提高徐紫薯8號(hào)作為粉條或類似產(chǎn)品加工原料的品質(zhì)。遮蔭處理下,T3時(shí)期徐紫薯8號(hào)鮮薯產(chǎn)量達(dá)最高且薯塊干物率較低,可作為鮮食型紫薯。此外,遮蔭可提高該時(shí)期可溶性糖含量,進(jìn)一步提升其鮮食品質(zhì)。就品質(zhì)而言,適當(dāng)早收可在保障徐紫薯8號(hào)鮮薯產(chǎn)量的前提下獲得塊根干物率較低的甘薯,食用品質(zhì)提升。夏季遮蔭后復(fù)光可提升徐紫薯8號(hào)作為粉絲、粉條等加工原料的品質(zhì)。因此,生產(chǎn)中應(yīng)考慮田間生長后期甘薯的產(chǎn)量、糖類物質(zhì)及淀粉特性的時(shí)期間差異,以及市場對(duì)不同用途甘薯塊根干物率、可溶性糖含量、淀粉含量和理化性質(zhì)的需求,優(yōu)化甘薯的收獲期。
4 結(jié)論
遮蔭導(dǎo)致煙薯25產(chǎn)量降低但會(huì)提升其食用品質(zhì),后期充足的光照可在保證其品質(zhì)的前提下彌補(bǔ)前期遮蔭造成的產(chǎn)量損失。徐紫薯8號(hào)對(duì)光環(huán)境具有較高的適應(yīng)性,在長期遮蔭和光環(huán)境變化下能維持產(chǎn)量穩(wěn)定,可在光能輻射資源較低的地區(qū)種植,作為鮮食甘薯時(shí)推薦適時(shí)早收,作為加工用途時(shí)適當(dāng)晚收可提高其淀粉加工品質(zhì)。
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(責(zé)任編輯 王暉)