盐胁迫对柳枝稷苗期生长和生理特性的影响

2010年,在人工气候室中设置了0、50、100、150和200 mmol·L-15种NaCl浓度处理,分析盐胁迫对柳枝稷苗期生长的影响.结果表明:随着NaCl浓度的增加,柳枝稷的生长明显受到抑制,株高降低、叶片变小、光合叶面积减少、净光合速率下降,干物质积累量显著降低,表现出甜土植物的特点.柳枝稷的耐盐能力较强,在200 mmol·L-1NaCl溶液中处理30 d后仍能存活,单株绿叶面积为491.9 cm2,净光合速率为0.93 μmol CO2·m-2·s-1.本试验条件下,以生长量下降50％为标准求得柳枝稷的耐盐阈值为178.6 mmol·L-1.     

硝酸镧浸种对NaCl胁迫下柳枝稷种子萌发及幼苗生理特性的影响

为探讨镧对NaCl胁迫下柳枝稷种子萌发及幼苗生理特性的影响,该研究以柳枝稷(Panicum virgatum L.)种子为研究材料,通过不同浓度(0、0.05、0.1、0.5和1.0mmol/L)La(NO3)3溶液浸种24h处理后,添加100mmol/L NaCl溶液,以蒸馏水为对照,在种子发芽箱中培养并观察柳枝稷种子萌发和幼苗生长,测定幼苗叶片相对电导率、丙二醛、脯氨酸和叶绿素的含量。结果显示:(1)0.05mmol/L La(NO3)3浸种处理可缓解100mmol/L NaCl对种子发芽的影响。(2)不同处理对幼苗苗长无显著影响,0.1mmol/L La(NO3)3浸种显著增加了幼苗的鲜重和叶面积,当浓度≥0.5mmol/L时根长、根冠比、鲜重和叶面积受到抑制。(3)幼苗叶片相对电导率和脯氨酸含量随La(NO3)3浓度的增加呈先降低后增加的趋势,其中0.05和0.1mmol/L La(NO3)3浸种处理下效果较好,丙二醛含量随La(NO3)3浓度的增加持续降低,但各浓度处理间差异不显著,叶绿素含量在0.05mmol/L La(NO3)3浸种处理下较NaCl处理显著增加,当浓度≥0.5mmol/L随着浸种浓度的增加呈下降趋势。(4)不同指标间发芽势与发芽率、根长和叶绿素含量间呈极显著正相关关系,与根冠比、电导率和脯氨酸含量均呈极显著负相关关系,而与苗长不相关。研究表明,低浓度(0.05mmol/L)La(NO3)3可缓解NaCl胁迫对柳枝稷种子萌发及幼苗生长的影响,而高浓度(1.0mmol/L)的La(NO3)3则会加重NaCl的胁迫危害。     

黄土丘陵区不同种植行距下柳枝稷光合生理日变化研究

以引进禾草柳枝稷为材料,在半干旱黄土丘陵区人工梯田设置20cm(L20)、40cm(L40)和60cm(L60)3种种植行距,比较研究了其光合生理参数日变化、地上部分生物量及土壤水分剖面分布特征,探讨其光合生理和水分利用特征与种植行距的关系。结果表明:(1)不同行距下,柳枝稷叶片净光合速率(Pn)日变化均呈双峰曲线,第一峰值均出现在10:00,第二峰值L40出现在14:00,L20和L60出现在16:00,具有明显的光合"午休"现象,且均由非气孔因素限制引起;柳枝稷叶片Pn日均值大小依次为:L60>L40>L20,且两两间差异显著。(2)柳枝稷叶片水分利用效率(WUE)日变化与Pn相似,分别在10:00和14:00达到峰值,日均值大小依次为:L60>L40>L20。(3)不同行距下,土壤体积含水量均随土层深度增加呈先增加后减少再增加的趋势,除0～50cm土层外,其它各土层土壤储水量均存在较大差异,0～380cm土层总土壤储水量表现为:L20>L40>L60。(4)柳枝稷地上生物量大小依次为:L20>L60>L40。研究发现,20cm行距的柳枝稷个体虽光合速率较低,但群体生物量和土壤储水量较高,所以柳枝稷在黄土丘陵区梯田的较优行距为20cm。     

柳枝稷长链非编码RNA PvLINC090588的功能研究

长链非编码RNA(long noncoding RNA, lncRNA)是一类长度大于200个核苷酸、无蛋白编码能力或蛋白编码能力较低的功能性RNA分子,可通过调控基因的表达影响植物的生长发育和对非生物胁迫的适应性。柳枝稷(Panicum virgatum L.)是一种抗旱性较强的禾本科(Poaceae)C₄生物能源模式作物。本研究从柳枝稷中克隆了1个新预测的lncRNA,将其命名为PvLINC090588。对该基因的表达模式分析显示：20%聚乙二醇、150 mmol/L氯化钠和50μmol/L脱落酸可诱导其表达,100μmol/L赤霉素抑制其表达,表明PvLINC090588的表达受多种非生物胁迫的诱导。过表达PvLINC090588转基因拟南芥(Arabidopsis thaliana)在100 mmol/L氯化钠、300 mmol/L甘露醇处理下其根长显著(P<0.05)大于野生型的,显示过表达株系的根系在胁迫下生长能力更强。与野生型相比,转基因株系在干旱和250 mmol/L氯化钠处理后其丙二醛含量降低,可溶性糖和脯氨酸含量显著升高(P<0.0...     

不同盐胁迫对柳枝稷生物量、品质和光合生理的影响

为明确不同盐胁迫对柳枝稷生物量、品质及光合生理的影响,以无盐胁迫作为对照(CK),选取0.40%Na Cl、0.80%Na2SO4和0.80%Na HCO3进行了土柱试验。结果表明:(1)与CK相比,Na Cl、Na2SO4、Na HCO3胁迫下柳枝稷地上生物量、地下生物量、总生物量、籽粒产量及根冠比均显著降低(P0.05),总生物量分别降低49.39%、60.52%、76.45%,Na HCO3对柳枝稷的生长抑制作用最强,Na Cl最弱;(2)Na Cl胁迫下柳枝稷地上生物质灰分含量显著增高14.89%,Na2SO4胁迫下硫(S)含量显著增高262.32%,纤维素含量显著降低13.71%,Na HCO3胁迫下钾(K)含量显著增高54.95%,半纤维素含量显著增高10.87%,灰分和S含量的增高不利于柳枝稷地上生物质的燃烧利用,纤维素含量的降低和半纤维素含量的增高不利于其转化利用;(3)Na Cl、Na2SO4、Na HCO3胁迫下柳枝稷叶片净光合速率(Pn)分别显著降低21.89%、29.54%和24.59%,气孔限制因素可能是其光合作用受到抑制、生物量下降的关键因素。     

利用SCoT分析柳枝稷遗传资源的多样性

本研究利用SCoT标记对96份柳枝稷种质的亲缘关系和遗传变异进行了研究。筛选出20条引物对96份供试材料进行PCR扩增,共获得445条带,其中多态性条带402条,平均多态性条带比率(PPB)达90.31%,多态性信息含量(PIC)为0.166~0.410,平均值为0.332,标记指数(MI)为10.20。遗传相似系数(GS)为0.498~0.912,平均值为0.688。表明SCoT标记能够揭示柳枝稷种质间的遗传变异。通过UPGMA分析表明,96份种质资源聚为高地型和低地型两大类。经POPGENE1.32软件分析结果显示:96份柳枝稷基因多样性指数(H)为0.285,Shannon指数(I)为0.431,表明供试的种质间遗传多样性丰富,遗传多样性水平高。经AMOVA 1.55方差分析揭示:96份柳枝稷生态型内的遗传变异占总变异的72.85%,生态型间遗传变异占总变异的27.15%,结果表明ScoT可用于柳枝稷遗传多样性研究,该研究结果可为柳枝稷种质资源的进一步开发利用提供重要信息。     

干旱胁迫下柳枝稷在露天矿区土壤中的种子萌发和生长特性

将能源植物用作矿区生态修复物种,对矿区的经济发展和生态环境具有重要意义。以能源植物柳枝稷为研究对象,通过盆栽试验,分析其在对照、轻度、中度以及重度干旱胁迫下柳枝稷幼苗在矿区土壤基质和非矿区土壤基质下的生长特性,并结合不同干旱胁迫下矿区土壤基质中种子萌发特征,揭示柳枝稷对干旱胁迫的响应机制和对矿区土壤的生态适应性。试验结果表明：(1)柳枝稷种子发芽总数、发芽率、发芽势以及发芽指数在轻度胁迫下达到最大值,活力指数在对照组最高,而在重度胁迫下,各项指标均达到最小值;种苗各生长指标在轻度胁迫下最小,幼苗根长和鲜重在重度胁迫下最高,芽长和芽重在其余三组胁迫下相差不大。(2)干旱胁迫使两种土壤基质下的柳枝稷株高降低,枯叶率增加,在对照、轻度胁迫和重度胁迫下能够保持叶片水分含量和分蘖数稳定;矿区土壤基质中,柳枝稷根体积在轻度胁迫下最大,中度胁迫下根长最长,重度胁迫下根数最多;根冠比随着干旱胁迫的加剧表现出先减后增的趋势,在轻度胁迫下最小,在重度胁迫下最大。(3)与非矿区土壤基质相比,柳枝稷在矿区土壤基质下根冠比更大,其余生长指标均更小;随着干旱胁迫的加剧,柳枝稷在矿区土壤基质下的株高、枯叶率、叶片相...     

黄土丘陵半干旱区引种禾草柳枝稷的生物量与水分利用效率

加强优良引种禾草植物的生态适应性研究对促进我国黄土高原半干旱区草地建设和草业科学发展具有十分重要的意义。比较研究了引种禾草柳枝稷(Panicumvirgatum)在黄土丘陵半干旱区不同立地条件下地上生物量的大小和季节累积差异及其水分利用特征。2001~2002年川地柳枝稷草地地上生物量达13000~16000kg/hm2,山地梯田和坡地为2300~2650kg/hm2。不同立地条件柳枝稷返青后的生物量累积过程呈二次或三次多项式。柳枝稷的绝对生长速率(AGR)在整个生长季内呈双峰曲线变化,川地柳枝稷草地的现存量和总量AGR最大值分别为158.93和169.83kg/(hm2·d),梯田分别为27.31和38.25kg/(hm2·d),坡地为37.0和36.69kg/(hm2·d)。坡地柳枝稷生物量较大值和AGR最大值出现时间最早。不同立地柳枝稷生物量相对生长速率(RGR)在整个生长季内呈双峰曲线变化,均以返青后的20d内最大,平均AGR以川地最大,坡地和梯田相近,但坡地两峰值高于梯田。不同立地柳枝稷草地土壤水分主要利用层次为0~2m,月平均含水量顺序为梯田>坡地>川地。川地柳枝稷叶片和整体生物量水分利用效率均最高,梯田整体生物量水分利用效率大于坡地,但二者叶片水分利用效率相近。川地和山地地表下5cm生育期平均地温均为17.60℃,但4~5月份川地地温高出山地1.2~2.8℃,川地4~10月份平均气温较山地高1.5℃,这些差异影响不同立地条件柳枝稷草地水分利用和生长进程。     

青贮对柳枝稷制取燃料乙醇转化过程的影响

青贮是一种传统的生物质原料保存方法,广泛应用于纤维素乙醇炼制领域尚需要考察其对原料品质和下游乙醇转化过程的影响。文中以秋季(初、中和末)收割的柳枝稷为原料,通过青贮、高温水热(LHW)预处理、纤维素酶水解和同步糖化与发酵(SSF)实验对上述问题予以回答。结果显示,秋季初收割的柳枝稷以不同湿度青贮后pH均小于4.0,干重损失小于2%,各主要成分与青贮前相比无明显变化;LHW预处理中青贮样品半纤维素水解率普遍高于未贮存样品,但青贮同样使原料获得了更高的发酵抑制物产生水平;青贮柳枝稷葡萄糖、木糖和半乳糖产量(预处理+酶水解)高于未贮存柳枝稷;经过168 h的SSF,青贮样品乙醇浓度为12.1 g/L,未贮存的秋季初、秋季中和秋季末柳枝稷为底物的浓度分别为10.3 g/L、9.7 g/L和10.6 g/L。综上,青贮有助于提高柳枝稷LHW预处理效率、酶水解率和乙醇产量。     

柳枝稷的生物学研究现状及其生物能源转化前景

柳枝稷(Panicum virgatum,L.)是起源于北美的一种热带草原高秆草,具有优良的草料特性和水土保持功能。因其有很大的能源利用前景,而目前全球能源紧缺,因此全世界范围内已有许多国家把研究柳枝稷转化生物能源这一利国利民的项目提到议事日程,展开了全方位的研究。这篇论文综述了柳枝稷的生物学研究现状以及其生物能源转化前景。     

Soil nitrogen dynamics in switchgrass seeded to a marginal cropland in South Dakota

The potential ecological impacts of switchgrass (Panicum virgatum L.), as a biofuel feedstock, have been assessed under different environmental conditions. However, limited information is available in understanding the integrated analysis of nitrogen (N) dynamics including soil nitrate (), nitrous oxide (N₂O) emissions, and leaching under switchgrass land management. The specific objective was to explore N dynamics for 2009 through 2015 in switchgrass seeded to a marginally yielding cropland based on treatments of N fertilization rate (N rate; low, 0; medium, 56; high, 112 kg N ha^?1) and landscape position (shoulder, backslope, and footslope). Our findings indicated that N rate impacted soil (0–5 cm depth) and surface N₂O fluxes but did not impact leaching during the observed years. Medium N (56 kg N ha^?1) was the optimal rate for increasing biomass yield with reduced environmental problems. Landscape position impacted the N dynamics. At the footslope position, soil , soil leaching, and N₂O fluxes were higher than the other landscape positions. Soil N₂O fluxes and leaching had downward trends over the observed years. Growing switchgrass on marginally yielding croplands can store soil N, reduce N losses via leaching, and mitigate N₂O emissions from soils to the atmosphere over the years. Switchgrass seeded on marginally yielding croplands can be beneficial in reducing N losses and can be grown as a sustainable bioenergy crop on these marginal lands.     

Simultaneous regulation of F5H in COMT‐RNAi transgenic switchgrass alters effects of COMT suppression on syringyl lignin biosynthesis

Ferulate 5‐hydroxylase (F5H) catalyses the hydroxylation of coniferyl alcohol and coniferaldehyde for the biosynthesis of syringyl (S) lignin in angiosperms. However, the coordinated effects of F5H with caffeic acid O‐methyltransferase (COMT) on the metabolic flux towards S units are largely unknown. We concomitantly regulated F5H expression in COMT‐down‐regulated transgenic switchgrass (Panicum virgatum L.) lines and studied the coordination of F5H and COMT in lignin biosynthesis. Down‐regulation of F5H in COMT‐RNAi transgenic switchgrass plants further impeded S lignin biosynthesis and, consequently, increased guaiacyl (G) units and reduced 5‐OH G units. Conversely, overexpression of F5H in COMT‐RNAi transgenic plants reduced G units and increased 5‐OH units, whereas the deficiency of S lignin biosynthesis was partially compensated or fully restored, depending on the extent of COMT down‐regulation in switchgrass. Moreover, simultaneous regulation of F5H and COMT expression had different effects on cell wall digestibility of switchgrass without biomass loss. Our results indicate that up‐regulation and down‐regulation of F5H expression, respectively, have antagonistic and synergistic effects on the reduction in S lignin resulting from COMT suppression. The coordinated effects between lignin genes should be taken into account in future studies aimed at cell wall bioengineering.     

Seed germination and seedling physiology of Larix kaempferi and Pinus densiflora in seedbeds with charcoal and elevated CO2

We investigated the effect of ectomycorrhizal colonization, charcoal and CO₂ levels on the germination of seeds of Larix kaempferi and Pinus densiflora, and also their subsequent physiological activity and growth. The seeds were sown in brown forest soil or brown forest soil mixed with charcoal, at ambient CO₂ (360 μmol mol⁻¹) or elevated CO₂ (720 μmol mol⁻¹), with or without ectomycorrhiza. The proportions of both conifer seeds that germinated in forest soil mixed with charcoal were significantly greater than for seeds sown in forest soil grown at each CO₂ level (P < 0.05; t-test). However, the ectomycorrhizal colonization rate of each species grown in brown forest soil mixed with charcoal was significantly lower than in forest soil at each CO₂ treatment [CO₂] (P < 0.01; t-test). The phosphorus concentrations in needles of each seedling colonized with ectomycorrhiza and grown in forest soil were greater than in nonectomycorrhizal seedlings at each CO₂ level, especially for L. kaempferi seedlings (P < 0.05; t-test), but the concentrations in seedlings grown in brown forest soil mixed with charcoal were not increased at any CO₂ level. Moreover, the maximum net photosynthetic rate of each seedling for light and CO₂ saturation (P _max) increased when the seedlings were grown with ectomycorrhiza at 720 μmol mol⁻¹ [CO₂]. Ectomycorrhizal colonization led to an increase in the stem diameter of each species grown in each soil treatment at each CO₂ level. However, charcoal slowed the initial growth of both species of seedling, constraining ectomycorrhizal development. These results indicate that charcoal strongly assists seed germination and physiological activity.     

Studies on the growth and development in Spirogyra

The phototropic response of Spirogyra sp. filaments and its relation with the different phases of their diurnal movements were studied. The filaments rapidly responded to unilateral irradiation; curvature towards the light source began in their tip region but shifted down to more basal regions. However, this typical and steady phototropic curvature could be observed only in the GnSt phase of the diurnal movement. In the other phases competitive states between the phototropic movement and other kinds of movement were evident. Thus, from the results of our previous and present studies it is proposed that the diurnal movement of Spirogyra filaments is composed of various individual movements, including a phototropic one; among these movements there exists a certain balance determined by the culture conditions and the time of day, and the phototropic movement is relatively inferior to the other movements.Abbreviations Gn negative gravitropism - Gp positive gravitropism - Sh shrinking (spiralling) growth - St straight form of growth V=Tanaka et al. (1983)     

煤粉尘沉降对鄂尔多斯高原优势植物羊柴幼苗生长的影响

随着鄂尔多斯煤矿开采的日益加剧,开采和运输过程中产生的煤粉尘沉降已成为影响当地植物生长的一个重要影响因子。通过近自然生境条件下的控制实验,以鄂尔多斯高原优势植物羊柴(Hedysarum laeve)为研究对象,探究不同梯度煤粉尘沉降量(0—3.5 mg/cm2)对其幼苗的光合生理特性和生长的影响。研究结果表明,随着煤粉尘沉降量的增加,羊柴叶片的气孔导度(Gs)、胞间二氧化碳浓度(Ci)、叶片水分饱和水汽压亏缺(Vpdl)等因子发生了不同程度的改变,导致净光合速率(Pn)和蒸腾速率(Tr)降低,使得幼苗的植株高度、叶片数、地上和地下生物量降低。因而,积累到一定程度的煤粉尘颗粒通过影响叶片气孔的水汽交换过程和降低叶片表面的光照强度,影响了叶片的光合生理过程,从而抑制了羊柴幼苗的生长。     

New evidence for a role for thioredoxin h in germination and seedling development

Thioredoxin of the h-type — earlier linked to the reduction of wheat (Triticum durum Desf. cv. Monroe) endosperm proteins — was converted from an oxidized to a partially reduced state during germination and seedling development. While the abundance of thioredoxin progressively decreased during this period, the availability of reducing equivalents, defined as the product of the relative abundance of thioredoxin and the percent reduction, increased. The amount of the enzyme catalyzing the reduction of thioredoxin h (NADP-thioredoxin reductase) remained constant. The activities of enzymes generating the NADPH needed for the reduction of thioredoxin (glucose 6-phosphate and 6-phosphogluconate dehydrogenases) increased. The level of thioredoxin h in the endosperm appeared to be controlled by the embryo via hormones. Gibberellic acid enhanced the disappearance of thioredoxin, whereas abscisic acid showed the opposite effect. Moreover, uniconazole, an inhibitor of gibberellic acid synthesis, slowed seedling growth and inhibited the disappearance of thioredoxin in a manner reversible by gibberellic acid. The results are consistent with a role for thioredoxin h in initiating the mobilization of nitrogen and carbon needed for germination and seedling development.Abbreviations ABA cis-abscisic acid - DTT dithiothreitol - GA gibberellin - GA₃ gibberellic acid - mBBr monobromobimane - NTR NADP-thioredoxin reductase This work was supported by National Science Foundation grant MCB-9316496. We thank Dr. Yuji Kamiya for advice and the Sumitomo Chemical Co. for a generous sample of uniconazole.     

Crop growth and development effects on surface albedo for maize and cowpea fields in Ghana,West Africa

The albedo () of vegetated land surfaces is a key regulatory factor in atmospheric circulation and plays an important role in mechanistic accounting of many ecological processes. This paper examines the influence of the phenological stages of maize (Zea mays) and cowpea (Vigna unguiculata) fields on observed albedo at a tropical site in Ghana. The crops were studied for the first and second planting dates in the year 2002. Crop management was similar for both seasons and measurements were taken from 10 m×10-m plots within crop fields. Four phenological stages were distinguished: (1) emergence, (2) vegetative, (3) flowering, and (4) maturity. measured from two reference surfaces, short grass and bare soil, were used to study the change over the growing seasons. Surface was measured and simulated at sun angles of 15, 30, 45, 60, and 75°. Leaf area index (LAI) and crop height (CH) were also monitored. Generally, increases from emergence to maturity for both planting dates in the maize field but slightly decreases after flowering in the cowpea field. For maize, the correlation coefficient (R) between and LAI equals 0.970, and the R between and CH equals 0.969. Similarly, for cowpea these Rs are 0.988 and 0.943, respectively. A modified albedo model adequately predicted the observed s with an overall R>0.860. The relative difference in surface with respect to the values measured from the two reference surfaces is discussed. Data presented are expected to be a valuable input in agricultural water management, crop production models, eco-hydrological models and in the study of climate effects of agricultural production, and for the parameterization of land-surface schemes in regional weather and climate models.     

水培条件下营养元素对枳幼苗根毛发育及根生长的影响

以柑橘砧木枳实生苗为试材,研究水培条件下N、P、K、Ca、Mg、Fe和Mn等7种营养元素分别缺乏对其根系主根长度、侧根数和主、侧根根毛密度、根毛长度及根毛直径等的影响.结果表明:水培条件下,不同缺素处理枳实生幼苗的根毛均能生长,但根毛主要集中在近根基段,根尖处分布较少;侧根的根毛密度显著大于主根,而其根毛长度显著小于主根.不同缺素处理对根毛的生长发育影响较大,主根根毛密度为55.0～174.3条·mm-2.与对照相比,缺Ca诱发主根的根毛密度、长度显著增加;缺P使主根的根基段、中段及侧根的根毛密度、长度显著增加;缺Fe使主根根尖段根毛密度显著增加,而长度显著降低;缺K使主根、侧根的根毛密度、长度及根毛直径均显著降低;缺Mg使主根根毛长度显著增加.各处理主根的生长较一致;侧根除缺N、Mg处理外,其他处理均出现脱落后再生的现象.     

蓝桉凋落叶分解对菠菜生长及光合特性的影响

通过盆栽试验,模拟蓝桉(Eucalyptus globulus)凋落叶在土壤中分解对受体植物菠菜(Spinacia oleracea)生长及光合特性的影响,设置L30(30 g/盆)、L60(60 g/盆)、L90(90 g/盆)和L120(120 g/盆)4个凋落叶施用水平,对照(CK)不加凋落叶。结果表明:(1)在分解30 d时,低量的凋落叶(L30)未抑制菠菜生物量的积累,而中量和高量处理(L60、L90和L120)下其生物量显著降低(P0.05);在凋落叶分解50 d后,蓝桉凋落叶的化感抑制效应已经减弱,仅L120仍然抑制菠菜生物量的积累;(2)一定量的凋落叶(L30和L60)对菠菜叶绿素的合成有促进作用,而凋落叶超过一定量时(L120)转为抑制作用,在凋落叶分解28 d时相对不明显,40 d后逐渐显现;(3)经凋落叶处理的菠菜叶片胞间CO2浓度(Ci)、气孔导度(Gs)及蒸腾速率(Tr)均显著高于对照(CK)(P0.05),而净光合速率(Pn)在L30处理下最高,L60和L90处理与CK差异不显著,仅L120处理显著低于CK(P0.05);(4)光响应与CO2响应曲线的特征参数表观量子效率(AQY)、最大净光合速率(Pmax)、光饱和点(Lsp)、光补偿点(Lcp)、暗呼吸速率(Rd)、Ru BP表观羧化效率(CE)和光呼吸速率(Rp)在L30处理下高于CK,其余凋落叶处理均低于CK,而CO2饱和点(Csp)、CO2补偿点(Ccp)随凋落叶量的增加而升高。综合各指标可见,少量的蓝桉凋落叶分解(L30)可通过增加叶绿素含量,提高对光和CO2的利用能力促进菠菜的光合作用,进而促进其生长,而凋落叶超过一定量时(L90—L120)则起到相反的作用。