徐佳楠 葛晨輝 王全華 王小麗
摘 要: 從營(yíng)養(yǎng)液配方、環(huán)境因素、水培馴化等方面綜述了有關(guān)菠菜水培的研究進(jìn)展.菠菜水培營(yíng)養(yǎng)液配方的研究主要集中在配方的篩選與主要營(yíng)養(yǎng)元素的調(diào)整熵.光照、溶解氧、液溫是影響水培菠菜生長(zhǎng)的重要環(huán)境因素.菠菜自身水培適應(yīng)性可能也是影響菠菜水培的重要因素,但目前鮮有相關(guān)報(bào)道.最后對(duì)今后的菠菜水培技術(shù)研究方向提出展望.
關(guān)鍵詞: 菠菜(Spinacia oleracea L.); 水培技術(shù); 營(yíng)養(yǎng)液配方; 環(huán)境因素; 水培適應(yīng)性
中圖分類(lèi)號(hào): S 636.1 ?文獻(xiàn)標(biāo)志碼: A ?文章編號(hào): 1000-5137(2019)05-0597-08
Abstract: This paper reviewed the research progress on leafy vegetable hydroponics about nutrient solution,environmental factors,cultivation measures and hydroponic domestication.The researches of spinach nutrient solution are mainly focused on the screening of nutrient solution formula and adjustment of main nutrient elements.Light,dissolved oxygen concentration and liquid temperature are main environmental factors influencing spinach growth under hydroponic conditions.Few researches have been conducted on the hydroponic adaptability of spinach.At last,the future research work on spinach hydroponics was also discussed.
Key words: Spinacia oleracea L.; hydroponic technology; nutrient solution; environmental factor; hydroponic adaptability
0 引 言
隨著都市農(nóng)業(yè)結(jié)構(gòu)的調(diào)整和消費(fèi)者對(duì)蔬菜質(zhì)量及食用安全意識(shí)的增強(qiáng),蔬菜水培技術(shù)因其在縮短生長(zhǎng)周期、減少水肥藥施用、增產(chǎn)保質(zhì)等方面的明顯優(yōu)勢(shì),日益成為國(guó)內(nèi)外設(shè)施園藝發(fā)展的重要技術(shù).蔬菜水培技術(shù)既適應(yīng)大都市對(duì)蔬菜產(chǎn)量和品質(zhì)的高需求,同時(shí)也是都市現(xiàn)代農(nóng)業(yè)科技水平的集中體現(xiàn).然而,目前我國(guó)水培蔬菜種類(lèi)主要以生菜類(lèi)、白菜類(lèi)為主.種類(lèi)單一限制了蔬菜水培技術(shù)的應(yīng)用范圍,因此有必要對(duì)其他蔬菜品種進(jìn)行水培專(zhuān)用新品種的選育及配套栽培技術(shù)的研究,以豐富水培蔬菜種類(lèi),促進(jìn)水培蔬菜的產(chǎn)業(yè)化發(fā)展.
菠菜(Spinacia oleracea L.)是我國(guó)普遍栽培的重要綠葉蔬菜之一,其營(yíng)養(yǎng)豐富,富含多種維生素,生產(chǎn)周期短,復(fù)種指數(shù)高,產(chǎn)量、產(chǎn)值高,是我國(guó)主要出口蔬菜之一.然而與其他葉菜類(lèi)蔬菜相比,目前市場(chǎng)上的菠菜品種大多是在傳統(tǒng)土培和基質(zhì)栽培的基礎(chǔ)上篩選培育而得,而在水環(huán)境,尤其是深液流水培系統(tǒng)中緩苗慢、生物產(chǎn)量低,無(wú)法展示其優(yōu)良性狀,嚴(yán)重限制了水培菠菜產(chǎn)量和質(zhì)量的提高.目前國(guó)內(nèi)對(duì)菠菜水培的研究主要集中在對(duì)已有營(yíng)養(yǎng)配方的改良上,但改善效果不明顯或適用面較窄[1].在篩選和培育水培專(zhuān)用菠菜新品種方面,更是鮮有研究報(bào)道.因此有必要總結(jié)相關(guān)葉菜水培生產(chǎn)經(jīng)驗(yàn),開(kāi)展水培條件下菠菜生命活動(dòng)規(guī)律的研究,為培育高產(chǎn)優(yōu)質(zhì)水培菠菜新品種,及提高菠菜水培生產(chǎn)技術(shù)提供依據(jù).
1 菠菜水培營(yíng)養(yǎng)液研究
1.1 營(yíng)養(yǎng)液配方比較
目前國(guó)內(nèi)菠菜水培采用的營(yíng)養(yǎng)液配方通常為各類(lèi)葉菜通用配方,如日本園試、日本山崎等.為篩選最適宜菠菜生長(zhǎng)的水培營(yíng)養(yǎng)液配方,王瑞等[2]比較了0.8倍Hoagland、日本山崎、日本園試、華南農(nóng)大葉菜A等4個(gè)葉菜類(lèi)常用營(yíng)養(yǎng)液配方對(duì)菠菜品質(zhì)、生物量動(dòng)態(tài)的影響,采用主成分分析法對(duì)生長(zhǎng)、品質(zhì)等多指標(biāo)進(jìn)行綜合評(píng)價(jià),其中,日本園試配方得分最高,是4種配方中最適宜菠菜水培的營(yíng)養(yǎng)液配方.本課題組比較了同一供氮水平(氮物質(zhì)的量濃度為8 mmol·L-1)的6個(gè)營(yíng)養(yǎng)液(0.76倍改良Hoagland、0.46倍日本園試、1.23倍日本山崎、0.38倍園藝均衡、華南農(nóng)大葉菜A、華南農(nóng)大葉菜B)對(duì)菠菜生長(zhǎng)的影響,結(jié)果發(fā)現(xiàn)人工氣候室靜態(tài)水培條件下,用1.23倍日本山崎配方營(yíng)養(yǎng)液培養(yǎng)的菠菜生物量、株高最高,而同期溫室淺液流水培條件下,用1.23倍日本山崎和0.38倍園藝均衡培養(yǎng)的菠菜生物量最高(該成果已被接受,待發(fā)表).在各營(yíng)養(yǎng)液總離子濃度比例保持不變的情況下,導(dǎo)致結(jié)果差異的原因可能與營(yíng)養(yǎng)液中營(yíng)養(yǎng)素絕對(duì)濃度有關(guān),尤其是硝態(tài)氮濃度.此外,水培方式以及栽培環(huán)境條件的差異,如不同水質(zhì)、葉菜種類(lèi)、生長(zhǎng)階段等也會(huì)影響營(yíng)養(yǎng)液配方的適用范圍.
1.2 營(yíng)養(yǎng)液中營(yíng)養(yǎng)元素的調(diào)節(jié)
為了優(yōu)化通用配方,很多研究者對(duì)營(yíng)養(yǎng)液中重要元素的濃度和配比對(duì)菠菜產(chǎn)量和品質(zhì)的影響開(kāi)展了大量研究,其中關(guān)于氮(N)、磷(P)、鉀(K)的研究報(bào)道較多,尤其是氮元素.水培營(yíng)養(yǎng)液氮素來(lái)源主要為硝態(tài)氮和銨離子,尤其是硝態(tài)氮.孫興祥等[3]以改進(jìn)的大澤營(yíng)養(yǎng)液為水培配方,研究了不同氮素水平(5,10,15,20 mmol·L-1)對(duì)4個(gè)菠菜品種生物量以及植株不同部位硝酸鹽、可溶性糖含量的影響,發(fā)現(xiàn)15 mmol·L-1氮素水平下菠菜生物量最高,菠菜的硝酸鹽含量隨氮素水平的提高呈遞增趨勢(shì),而可溶性糖含量則隨氮素水平的提高呈遞減趨勢(shì).OKAZAKI等[4]比較了3個(gè)硝態(tài)氮濃度水平(1,2,4 mmol·L-1)對(duì)水培菠菜的影響,發(fā)現(xiàn)4 mmol·L-1硝態(tài)氮濃度下菠菜的產(chǎn)量、總氮含量和硝酸鹽含量均最高.作者還設(shè)置了不同物質(zhì)的量濃度比例的硝態(tài)氮/銨態(tài)氮(10 ?∶ 0,5 ?∶ 5,3 ?∶ 7)的營(yíng)養(yǎng)液配方處理,發(fā)現(xiàn)3個(gè)處理對(duì)水培菠菜鮮重、總氮含量影響無(wú)顯著差異,但隨著銨態(tài)氮比例的增加,植株硝態(tài)氮含量明顯降低.其他研究報(bào)道也發(fā)現(xiàn)類(lèi)似的規(guī)律,當(dāng)NH4+-N/NO3--N物質(zhì)的量濃度比值從100 ?∶ 0變化到0 ?∶ 100,菠菜的生物量、株高、根系長(zhǎng)度、硝酸鹽和亞硝酸鹽累積量等均呈增加趨勢(shì),營(yíng)養(yǎng)液中適當(dāng)增銨能提高菠菜品質(zhì),但不能增加菠菜的產(chǎn)量.NH4+-N/NO3--N比值為0 ?∶ 100時(shí),菠菜莖葉生物量最高,隨著NH4+-N/NO3--N 比例的增加,菠菜的生物量和硝酸鹽累積量均呈遞減趨勢(shì)[5-6],水溶性糖含量則呈遞增趨勢(shì)[6].除硝態(tài)氮和銨態(tài)氮外,適當(dāng)配施有機(jī)氮能顯著提高營(yíng)養(yǎng)效應(yīng),改善作物品質(zhì).不同物質(zhì)的量濃度比例的硝態(tài)氮/甘氨酸處理(100 ?∶ 0,75 ?∶ 25,50 ?∶ 50,25 ?∶ 75,0 ?∶ 100)下,甘氨酸部分替代硝態(tài)氮降低了菠菜生物量,但同時(shí)硝態(tài)氮含量也顯著降低[7].
2.3 溫 度
溫度是影響水培植物生長(zhǎng)的重要因素之一.GENT[32]研究發(fā)現(xiàn)生菜相對(duì)生長(zhǎng)速度(RGR)更容易受日照輻射的影響,而受溫度和營(yíng)養(yǎng)液硝態(tài)氮濃度影響不大;相反,菠菜更容易受溫度和營(yíng)養(yǎng)液硝態(tài)氮濃度影響.SEO等[31]也發(fā)現(xiàn)不管營(yíng)養(yǎng)液的DOC濃度如何,夏季高溫是導(dǎo)致水培菠菜死亡的直接原因,而生菜不受營(yíng)養(yǎng)液溫度的影響.但沒(méi)有關(guān)于菠菜水培適宜溫度范圍的詳細(xì)報(bào)道.
3 菠菜自身因素及其他
菠菜自身對(duì)水培環(huán)境的適應(yīng)能力是影響其水培生產(chǎn)表現(xiàn)的關(guān)鍵因素之一.不同作物對(duì)環(huán)境條件的適應(yīng)性不同,同一作物不同品種對(duì)水培環(huán)境的適應(yīng)性也有差異.SINHA等[33]發(fā)現(xiàn)小麥品種間對(duì)不同栽培介質(zhì)(水培、基質(zhì)、土培)的響應(yīng)存在較大差異,說(shuō)明可以通過(guò)品種篩選獲得水培適應(yīng)性能力高的作物品種.根系是影響植物水培適應(yīng)性的關(guān)鍵器官.水培環(huán)境下,根系環(huán)境發(fā)生很大的變化,根系環(huán)境的變化又會(huì)顯著影響植物對(duì)水分和養(yǎng)分的吸收,進(jìn)而影響植物的生長(zhǎng)[34].如何利用菠菜各器官尤其是根系的適應(yīng)性反應(yīng)以充分挖掘菠菜自身對(duì)水環(huán)境的適應(yīng)潛力,是提高菠菜水培生產(chǎn)效率的關(guān)鍵之一.已有研究報(bào)道水培環(huán)境下植物能夠改變地上部分與根系的形態(tài),及微觀結(jié)構(gòu)以適應(yīng)低氧的水培環(huán)境.如耐水培的植物,其水環(huán)境中的植物根皮層內(nèi)一般會(huì)形成發(fā)達(dá)的通氣組織,以利于根系供氧[35].水培低氧環(huán)境還會(huì)誘導(dǎo)不定根、根中徑向氧氣損失屏障的形成,以及莖基或下胚軸肥腫等形態(tài)結(jié)構(gòu)的變化.這些變化均有利于地上部的氧氣向根系擴(kuò)散,緩解低氧對(duì)植物的脅迫作用[36-37].目前為止在菠菜水培適應(yīng)性機(jī)理方面的研究未見(jiàn)報(bào)道,后續(xù)可以從菠菜根系微觀結(jié)構(gòu)、生理生化及分子機(jī)制方面開(kāi)展相關(guān)研究.
還有研究發(fā)現(xiàn),水培體系由于缺乏培養(yǎng)介質(zhì)對(duì)根際分泌物的吸附、中和,在長(zhǎng)期水培過(guò)程中,蔬菜根系分泌出的毒性物質(zhì)經(jīng)累積后會(huì)造成自毒危害,抑制蔬菜生長(zhǎng)[38].常見(jiàn)的各類(lèi)蔬菜如豌豆[39]、生菜[40]、黃瓜[41-42]、西紅柿[43]、蘆筍[44]等都可釋放自毒物質(zhì),采用活性炭吸附、光催化法等方法可去除植物自毒物質(zhì),減輕其對(duì)植物的生長(zhǎng)抑制作用[38].菠菜根際分泌物是否會(huì)對(duì)水培菠菜的生長(zhǎng)產(chǎn)生抑制作用還未見(jiàn)報(bào)道.
4 展 望
菠菜是重要的生、熟食綠葉蔬菜,在水培生產(chǎn)中具有廣闊的應(yīng)用前景.開(kāi)發(fā)適合多種水培生產(chǎn)模式的菠菜水培生產(chǎn)技術(shù),培育水培專(zhuān)用高產(chǎn)優(yōu)質(zhì)菠菜新品種是實(shí)現(xiàn)菠菜水培規(guī)?;a(chǎn)的前提.然而與水培生菜相比,目前菠菜水培技術(shù)研究基礎(chǔ)薄弱,無(wú)論是關(guān)于營(yíng)養(yǎng)液配方還是環(huán)境因素調(diào)控方面的報(bào)道均較少,在菠菜水培專(zhuān)用品種的選育方面更是匱乏.今后有必要從以下幾個(gè)方面重點(diǎn)開(kāi)展菠菜水培技術(shù)相關(guān)的基礎(chǔ)研究:1) 繼續(xù)開(kāi)展對(duì)菠菜水培營(yíng)養(yǎng)液關(guān)鍵營(yíng)養(yǎng)元素最適濃度的研究,明確菠菜不同生育期對(duì)各營(yíng)養(yǎng)元素需求的動(dòng)態(tài)變化,尤其是N,P與K的絕對(duì)濃度與濃度配比;2) 考慮環(huán)境因素相互之間可能存在耦合效應(yīng),開(kāi)展菠菜水培環(huán)境的多因素綜合調(diào)控研究,探索最優(yōu)的綜合環(huán)境調(diào)控技術(shù)參數(shù);3) 深入研究菠菜根系應(yīng)對(duì)液溫變化與低氧脅迫的響應(yīng)機(jī)制,與綜合調(diào)控策略;4) 參考其他植物水生馴化機(jī)制,開(kāi)展菠菜水培馴化相關(guān)的形態(tài)、生理生化特征研究,建立菠菜耐水培品種評(píng)價(jià)指標(biāo)體系,篩選耐水培菠菜種質(zhì)資源,并在此基礎(chǔ)上進(jìn)一步探究菠菜水培馴化機(jī)制,挖掘菠菜根系水生相關(guān)基因.
參考文獻(xiàn):
[1] 王瑞,胡笑濤,蘇苑君.菠菜營(yíng)養(yǎng)液栽培的研究進(jìn)展與展望 [J].長(zhǎng)江蔬菜,2014(14):7-10.
WANG R,HU X T,SU Y J.Research progress and prospect on nutrient solution cultivation of spinach [J].Journal of Changjiang Vegetables,2014(14):7-10.
[2] 王瑞,胡笑濤,王文娥,等.水培菠菜不同配方的產(chǎn)量、品質(zhì)主成分分析研究 [J].北方園藝,2016(10):27-31.
WANG R,HU X T,WANG W E,et al.Study on different nutrient solution formula hydroponics on spinach yield,quality and major elements of consumption [J].Northern Horticulture,2016(10):27-31.
[3] 孫興祥,王健,周毅,等.不同氮素水平對(duì)菠菜生長(zhǎng)和品質(zhì)的影響 [J].南京農(nóng)業(yè)大學(xué)學(xué)報(bào),2005(3):126-128.
SUN X X,WANG J,ZHOU Y,et al.Effects of different nitrogen levels on the growth and quality of spinach [J].Journal of Nanjing Agricultural University,2005(3):126-128.
[4] OKAZAKI K,OKA N,SHINANO T,et al.Differences in the metabolite profiles of spinach (Spinacia oleracea L.) leaf in different concentrations of nitrate in the culture solution [J].Plant and Cell Physiology,2008,49(2):170-177.
[5] 邢素芝,汪建飛,李孝良,等.氮肥形態(tài)及配比對(duì)菠菜生長(zhǎng)和安全品質(zhì)的影響 [J].植物營(yíng)養(yǎng)與肥料學(xué)報(bào),2015,21(2):527-534.
XING S Z,WANG J F,LI X L,et al.Different nitrogen fertilizers and ratios effect on growth,safety and quality of spinach [J].Journal of Plant Nutrition and Fertilizer,2015,21(2):527-534.
[6] 王健,孫興祥,沈其榮,等.增銨對(duì)菠菜生長(zhǎng)及品質(zhì)的影響 [J].土壤通報(bào),2006(2):2326-2329.
WANG J,SUN X X,SHEN Q R,et al.Effects of enhanced ammonium nutrition on the growth and quality of spinach [J].Chinese Journal of Soil Science,2006(2):2326-2329.
[7] LIU X,WANG L,LI Z,et al.Nitrate/Gly ratios in nutrition influenced the growth and amino acid composition in spinach (Spinacia oleracea L.) [J].Journal of Plant Nutrition,2013,37(5):765-776.
[8] 蘇苑君,胡笑濤,王文娥,等.磷對(duì)水培生菜生長(zhǎng)及礦質(zhì)元素動(dòng)態(tài)吸收的影響 [J].中國(guó)生態(tài)農(nóng)業(yè)學(xué)報(bào),2015,23(10):1244-1252.
SU Y J,HU X T,WANG W Z,et al.Effect of phosphorus on dynamic growth and nutrient absorption of hydroponic [J].Chinese Journal of Eco-Agriculture,2015,23(10):1244-1252.
[9] 汪建飛,董彩霞,謝越,等.銨硝比和磷素營(yíng)養(yǎng)對(duì)菠菜生長(zhǎng)、氮素吸收和相關(guān)酶活性的影響 [J].土壤學(xué)報(bào),2006(6):954-960.
WANG J F,DONG C X,XIE Y,et al.Effects of NH4+-N/ NO3--N ratio and phosphorus levels on growth,nitrogen uptake and relevant enzymes activity of spinach (Spinacia oleracea L.) [J].Acta Pedologica Sinica,2006(6):954-960.
[10] 陶其驤,羅奇祥,劉光榮,等.施鉀對(duì)改善作物產(chǎn)品品質(zhì)的效果 [J].江西農(nóng)業(yè)學(xué)報(bào),1999,11(3):29-34.
TAO Q X,LUO Q X,LIU G R,et al.Effect of K application on quality of crop products [J].Acta Agriculture Jiangxi,1999,11(3):29-34.
[11] 孫紅梅,李天來(lái),須暉,等.不同氮水平下鉀營(yíng)養(yǎng)對(duì)大棚番茄產(chǎn)量及品質(zhì)的影響 [J].沈陽(yáng)農(nóng)業(yè)大學(xué)學(xué)報(bào),2000,31(1):68-71.
SUN H M,LI T L,XU H,et al.Effects of potassium fertilizers on yield and quality of tomato under differrent application of nitrogen fertilizers [J].Journal of Shenyang Agricultural University,2000,31(1):68-71.
[12] 蘇苑君,胡笑濤,王文娥,等.鉀濃度對(duì)水培生菜生長(zhǎng)及礦質(zhì)元素動(dòng)態(tài)吸收的影響 [J].西北農(nóng)林科技大學(xué)學(xué)報(bào)(自然科學(xué)版),2016,44(8):191-196,204.
SU Y J,HU X T,WANG W E,et al.Effect of potassium concentration on growth and dynamic absorption of mineral elements of hydroponic lettuce [J].Journal of Northwest A&F University (Natural Science Edition),2016,44(8):191-196,204.
[13] 方素萍.氮鉀營(yíng)養(yǎng)對(duì)菠菜生長(zhǎng)、硝酸鹽累積的影響及機(jī)理研究 [D].杭州:浙江大學(xué)2002.
FANG S P.Nitrate accumulation in spinach (Spinacia oleracea L.) [D].Hangzhou:Zhejiang University,2002.
[14] 于洪波.氮鉀營(yíng)養(yǎng)對(duì)蔬菜累積草酸的調(diào)控及其機(jī)理研究 [D].杭州:浙江大學(xué),2002.
YU H B.Manipulation of oxalate accumulation in vegetable crops through Nitrogen (N) and Potassium (K) [D].Hangzhou:Zhejiang University,2002.
[15] NIU G H,SUN Y P,MASABNI J G.Impact of low and moderate salinity water on plant performance of leafy vegetables in a recirculating NFT System [J].Horticulturae,2018,4(1):6.
[16] JIN C W,LIU Y,MAO sQ Q,et al.Mild Fe-deficiency improves biomass production and quality of hydroponic-cultivated spinach plants (Spinacia oleracea L.) [J].Food Chemistry,2013,138(4):2188-2194.
[17] 李登超,朱祝軍,徐志豪.硒對(duì)菠菜抗氧化系統(tǒng)及過(guò)氧化氫含量的影響 [J].園藝學(xué)報(bào),2002(6):547-550.
LI D C,ZHU Z J,XU Z H.Effects of selenium on antioxidative system and H2O2 content in spinach [J].Acta Horticulturae Sinica,2002(6):547-550.
[18] 王魏,鄒志榮,喬飛,等.外源ALA對(duì)NaCl脅迫下菠菜生理特性的影響 [J].西北農(nóng)業(yè)學(xué)報(bào),2008(1):137-141,156.
WANG W,ZOU Z R,QIAO F,et al.Effects of exogenous ALA on physiological characteristics of spinach under NaCl stress [J].Acta Agriculturae Boreali-Occidentalis Sinica,2008(1):137-141,156.
[19] 王穎,郭世榮,束勝,等.外源亞精胺對(duì)鹽脅迫下菠菜葉綠素合成前體含量的影響 [J].西北植物學(xué)報(bào),2015(10):2026-2034.
WANG Y,GUO S R,SHU S,et al.Effects of exogenous spermidine on chlorophyll precursors content of spinach plants under salt stress [J].Acta Botanica Boreali-Occidentalia Sinica,2015(10):2026-2034.
[20] 陳冠男,劉金香,曹宇,等.水楊酸對(duì)鹽脅迫下菠菜葉表皮氣孔開(kāi)度的調(diào)節(jié) [J].鄭州輕工業(yè)學(xué)院學(xué)報(bào)(自然科學(xué)版),2014(6):33-38.
CHEN G N,LIU J X,CAO Y,et al.The adjustment of salicylic acid on stomatal aperture of Spinacia oleracea L.leaf epidermis under salt stress [J].Journal of Zhengzhou University of Light Industry (Natural Science Edition),2014(6):33-38.
[21] ZEEVAART J A,GAGE D A,TALON M.Gibberellin A1 is required for stem elongation in spinach [J].Proceedings of the National Academy of Sciences of the United States of America,1993,90(15):7401-7405.
[22] 羅樹(shù)生,胡華敏,王宗抗,等.植物生長(zhǎng)調(diào)節(jié)劑灌根對(duì)菠菜的作用效果研究 [J].廣東農(nóng)業(yè)科學(xué),2012,39(19):64-66,70.
LUO S S,HU H M,WANG Z K,et al.Regulation effects of plant growth regulators on Spinacia oleracea L.by soil drench application [J].Guangdong Agricultural Sciences,2012,39(19):64-66,70.
[23] 王燦,蘇艷,李樹(shù),等.不同光照強(qiáng)度對(duì)室內(nèi)水培葉用蔬菜生長(zhǎng)的影響 [J].天津農(nóng)學(xué)院學(xué)報(bào),2014,21(2):18-20,23.
WANG C,SU Y,LI S,et al.Effects of different light intensity on indoor hydroponic leafy vegetable growth [J].Journal of Tianjin Agricultural University,2014,21(2):18-20,23.
[24] PROIETTI S,MOSCATELLO S,LECCESE A,et al.The effect of growing spinach (Spinacia oleracea L.) at two light intensities on the amounts of oxalate,ascorbate and nitrate in their leaves [J].Journal of Horticultural Science and Biotechnology,2004,79(4):606-609.
[25] YORIO N C,GOINS G D,KAGIE H R,et al.Improving spinach,radish,and lettuce growth under red light-emitting diodes (LEDs) with blue light supplementation [J].HortScience,2001,36(2):380-383.
[26] OHASHI-KANEKO K,TAKASE M,KON N,et al.Effect of light quality on growth and vegetable quality in leaf lettuce,spinach and komatsuna [J].Environment Control in Biology,2007,45(3):189-198.
[27] 黃碧陽(yáng),林碧英,李彩霞,等.LED光質(zhì)對(duì)菠菜生長(zhǎng)和光合生理特性的影響 [J].福建農(nóng)林大學(xué)學(xué)報(bào)(自然科學(xué)版),2018,47(4):403-408.
HUANG B Y,LIN B Y,LI C X,et al.Effects of LED light quality on growth and photosynthetic physiological characteristics in spinach [J].Journal of Fujian Agriculture and Forestry University (Natural Science Edition),2018,47(4):403-408.
[28] 黃碧陽(yáng),林碧英,李彩霞,等.LED紅藍(lán)光配比對(duì)菠菜生長(zhǎng)及品質(zhì)的影響 [J].江蘇農(nóng)業(yè)科學(xué),2018,46(7):131-135.
HUANG B Y,LIN B Y,LI C X,et al.Effects of red and blue ratios of LED on growth and quality of spinach [J].Jiangsu Agricultural Sciences,2018,46(7):131-135.
[29] 黃傳輝.菠菜生長(zhǎng)最適光環(huán)境的研究 [D].福州:福建農(nóng)林大學(xué),2016.
HUANG C H.The optimal light environment for spinach growth [D].Fuzhou:Fujian Agriculture and Forestry University,2016.
[30] CHUN C,TAKAKURA T.Rate of root respiration of lettuce under various dissolved oxygen concentrations in hydroponics [J].Environment Control in Biology,1994,32(2):125-135.
[31] SEO T C,KIM Y C,LEE J W,et al.The effect of dissolved oxygen concentration on the growth and nutrient uptake of spinach and lettuce grown hydroponically in summer season [J].Journal of the Korean Society for Horticultural Science,2002,43(4):421-424.
[32] GENT M P N.Factors affecting relative growth rate of lettuce and spinach in hydroponics in a greenhouse [J].Hortscience,2017,52(12):1742-1747.
[33] SINHA S K,RANI M,KUMAR A,et al.Natural variation in root system architecture in diverse wheat genotypes grown under different nitrate conditions and root growth media [J].Theoretical and Experimental Plant Physiology,2018,30(3):223-234.
[34] 牛曉麗.作物根系對(duì)局部供應(yīng)水氮的響應(yīng)及其生理機(jī)制 [D].楊凌:西北農(nóng)林科技大學(xué),2016.
NIU X L.Effects of partial supply of water and nitrogen on root growth and absorption capacity [D].Yangling:Northwest A&F University,2016.
[35] 樊明壽,張福鎖.植物通氣組織的形成過(guò)程和生理生態(tài)學(xué)意義 [J].植物生理學(xué)通訊,2002(6):615-618.
FAN M S,ZHANG F S.Aerenchyma formation in plant and its physiological and ecological significance [J].Plant Physiology Communications,2002(6):615-618.
[36] 馬月花,郭世榮,杜南山,等.低氧脅迫對(duì)黃瓜幼苗生長(zhǎng)和形態(tài)結(jié)構(gòu)及有關(guān)酶活性的影響 [J].南京農(nóng)業(yè)大學(xué)學(xué)報(bào),2016,39(2):213-219.
MA Y H,GUO S R,DU N S,et al.Effect of hypoxia stress on growth,morpho-anatomical acclimation and activity of involved enzymes of cucumber seedlings [J].Journal of Nanjing Agricultural University,2016,39(2):213-219.
[37] 汪天,王素平,郭世榮,等.植物低氧脅迫傷害與適應(yīng)機(jī)理的研究進(jìn)展 [J].西北植物學(xué)報(bào),2006(4):847-853.
WANG T,WANG S P,GUO S R,et al.Research advance about hypoxia-stress damage and hypoxia-stress-adapting mechanism in plants [J].Acta Botanica Boreali-Occidentalia Sinica,2006(4):847-853.
[38] 劉文科,楊其長(zhǎng).設(shè)施無(wú)土栽培營(yíng)養(yǎng)液中植物毒性物質(zhì)的去除方法 [J].北方園藝,2010(16):69-70.
LIU W K,YANG Q C.Remove of phytotoxic substances from nutrient solution of soilless cultureunder cover [J].Northern Horticulture,2010(16):69-70.
[39] 韓旭,杜公福,牛玉,等.水培法收集甜椒根系分泌物化學(xué)成分鑒定 [J].長(zhǎng)江蔬菜,2016(4):47-52.
HAN X,DU G F,NIU Y,et al.Chemical components identification on root exudates of Capsicum Fructescens L.by nutrient solution culture [J].Journal of Changjiang Vegetables,2016(4):47-52.
[40] LEE J G,LEE B Y,LEE H J.Accumulation of phytotoxic organic acids in reused nutrient solution during hydroponic cultivation of lettuce (Lactuca sativa L.) [J].Scientia Horticulturae,2006,110(2):119-128.
[41] 鄒麗蕓.西瓜連作障礙中自毒作用的研究 [D].杭州:浙江大學(xué),2004.
ZOU L Y.Study on autotoxicity in continuous cropping obstacle of watermelon plant [D].Hangzhou:Zhejiang University,2004.
[42] YU J Q,MATSUI Y.Phytotoxic substances in root exudates of cucumber (Cucumis sativus L.) [J].Journal of Chemical Ecology,1994,20(1):21-31.
[43] YU J Q,MATSUI Y.Extraction and identification of phytotoxic substances accumulated in nutrient solution for the hydroponic culture of tomato [J].Soil Science and Plant Nutrition,1993,39(4):691-700.
[44] SUNADA K,DING X G,UTAMI M S,et al.Detoxification of phytotoxic compounds by TiO2 photocatalysis in a recycling hydroponic cultivation system of asparagus [J].Journal of Agricultural and Food Chemistry,2008,56(12):4819-4824.
(責(zé)任編輯:顧浩然)