冷锻炼处理减轻低温贮藏桃果实冷害的能量代谢机理

为探讨通过维持低温贮藏桃果实能量代谢水平而减轻冷害的有效方法,以‘大久保’桃为试材,分别以24、5、0℃贮藏为对照,研究冷锻炼处理(8℃冷锻炼5d后转入0℃冷藏,CAT)对维持冷藏及其货架期间桃果实线粒体呼吸代谢相关酶及三磷酸腺苷(ATP)含量的影响。结果表明:24℃贮藏模式下桃果实的H+-ATPase、Ca2+-ATPase、琥珀酸脱氢酶(SDH)和细胞色素氧化酶(CCO)活性在贮藏过程中2～3d出现了活性高峰,之后随着果实的软化衰老而显著下降。5℃贮藏桃果实贮藏期间的H+-ATPase、Ca2+-ATPase、SDH、CCO活性和ATP含量最低,且下降速度也最快;0℃贮藏次之,而CAT贮藏可有效抑制桃果实H+-ATPase、SDH和CCO活性的下降,保持较高的ATP含量。随着贮藏时间的延长,货架期期间H+-ATPase、Ca2+-ATPase、SDH、CCO活性和ATP含量均降低,但CAT处理明显高于5、0℃冷藏处理。这表明冷锻炼处理可通过调节线粒体呼吸代谢酶活性,维持果实较高的能量水平,有利于减轻果实冷害的发生并延长贮藏期。     

外源亚精胺对盐胁迫下黄瓜幼苗活性氧代谢的影响

采用营养液水培,研究了盐胁迫下外源亚精胺(Spd)对抗盐能力不同的2个黄瓜品种幼苗叶片和根系内活性氧(ROS)水平和抗氧化酶活性的影响.结果表明,外源Spd降低了50 mmol/L NaCl胁迫下幼苗体内O-.2产生速率、H2O2和丙二醛(MDA)含量,提高了抗氧化酶SOD、POD和CAT的活性;与抗盐性较强的"长春密刺"品种相比,Spd处理对抗盐性较弱的"津春二号"品种效果更明显.表明外源Spd处理通过促进盐胁迫下植株体内抗氧化酶活性的提高,降低ROS水平,从而缓解NaCl对黄瓜幼苗的盐伤害.     

低温贮藏对桃果实冷害和能量水平的影响

为探索桃果实低温贮藏时冷害发生与能量水平的关系,该研究将桃果实分别置于0、5、10℃下贮藏,定期测定果实褐变指数、硬度和出汁率等冷害指标以及H+-ATPase、Ca2+-ATPase、琥珀酸脱氢酶(SDH)、细胞色素氧化酶(CCO)等线粒体呼吸代谢相关酶活性及能量相关物质三磷酸腺苷(ATP)、二磷酸腺苷(ADP)、磷酸腺苷(AMP)含量和能荷变化。结果表明,桃果实在不同贮藏温度下冷害发生程度不同,5℃贮藏的桃果实褐变指数最高,冷害最严重,0℃贮藏次之,10℃贮藏无冷害发生。同时发现5℃贮藏果实的ATP和ADP含量、能荷水平、H+-ATPase、SDH和CCO活性最低。这些结果表明,桃果实冷害发生可能与能量供应不足有关。     

葡萄果实转色过程中线粒体呼吸代谢变化

为明确线粒体呼吸代谢与果实成熟期间颜色形成的关系,本试验探究了3种葡萄果实(夏黑、巨峰和金手指)成熟期间颜色、总花色苷含量、呼吸强度、内源乙烯产生和线粒体呼吸代谢的变化规律。结果表明,夏黑和巨峰葡萄果实的果皮颜色指数(CIRG)值和总花色苷含量随着果实成熟呈上升趋势,而金手指葡萄果实CIRG值在整个成熟期间变化不显著;葡萄果实成熟期间呼吸强度和内源乙烯均呈先上升后下降趋势,在花后63d达到峰值。同时,葡萄果实成熟期间,线粒体膜通透性转换孔(MPTP)逐渐增加,线粒体膜流动性(MMF)持续下降;线粒体琥珀酸脱氢酶(SDH)活性随着成熟进程逐渐下降,而细胞色素氧化酶(CCO)活性在花后63d开始上升。表明盛花63d之后是果实色泽快速形成阶段,可能为葡萄果实色泽形成的关键时期。线粒体呼吸代谢增强促进了果实成熟期间呼吸强度增加和内源乙烯释放,从而使夏黑和巨峰果实中花色苷大量积累,加快了金手指葡萄果实叶绿素的降解和类胡萝素的合成。本研究结果为调控南方葡萄果实着色提供了理论依据。     

盐胁迫对紫苏叶片光合特性及离子平衡的影响

以紫苏实生苗为材料,以正常供水为对照,研究不同盐胁迫程度对紫苏叶片光合特性及离子平衡的影响,以期为紫苏规范化栽培提供依据。结果表明,在盐胁迫下,紫苏叶片中丙二醛(MDA)含量、相对电导率均随着NaCl浓度的升高而增加,光合参数(Pn、Gs、Ci、Tr)则均随着盐胁迫程度的加强而下降;光合色素和抗氧化酶活性随着NaCl浓度的增加均呈现出先增加后下降的趋势,光合色素含量则在NaCl浓度为25 mmol L-1时达到最大值,抗氧化酶(SOD、POD、CAT)活性则在Na Cl浓度为40 mmol L-1时达到最大值。紫苏叶片中Na+、K+、Ca2+含量在低盐浓度条件下变化缓慢,在高盐浓度下Na+含量显著增加而K+、Ca2+含量则显著下降。本试验结果说明紫苏能够忍耐一定程度的盐胁迫伤害,对盐有一定的适应性。     

不同浓度盐和H2O2对海马齿PHGPx活性的影响

磷脂氢谷胱甘肽过氧化物酶(PHGPx)是目前发现的唯一能够直接还原膜上脂类过氧化物的抗氧化酶,在保护生物膜免受过氧化损伤方面发挥重要作用。本研究探讨了海马齿PHGPx活性的测定,检测了不同浓度盐和 H2O2胁迫对PHGPx活性的影响。结果显示,以蒸馏水为缓冲液提取的叶片总蛋白效果较好; NaCl梯度浓度处理下,海马齿叶片PHGPx活性呈先降低后升高然后再降低的趋势,其中500mmol/L NaCl处理可以诱导最大活性; H2O2梯度浓度处理下,海马齿叶片PHGPx活性呈先升高后降低再升高趋势,0.5mmol/L H2O2处理获得最大活性;海马齿植株经 H2O2清除剂DMTU处理后再用 H2O2处理,PHGPx的活性降低,同时 NaCl的诱导效果并不受到影响。这些研究结果表明,海马齿中PHGPx的活性受到盐和 H2O2的调节,并且它们对PHGPx酶活的调节可能是两个独立的过程。     

复合微生物肥料对盐胁迫下番茄生理特性的影响

采用盆栽试验的方法,研究不同NaCl浓度(0、20 m M、50 m M和100 m M)胁迫下,添加复合微生物肥料对番茄幼苗生长、光合色素含量、根系活力和抗氧化指标的影响。结果显示,低盐胁迫(20 m M NaCl)促进了番茄幼苗的生长,但随盐胁迫浓度的升高,生物量逐渐降低,同时降低了根系活力同时下降;不同盐浓度胁迫均提高了番茄叶片内光合色素含量、根系和叶片内抗氧化酶(SOD、POD、CAT)活性、丙二醛(MDA)含量和超氧阴离子自由基(O2.-)产生速率。添加复合微生物肥料显著提高了盐胁迫番茄幼苗生物量、根系活力,、进一步提高了叶片内光合色素含量和根系抗氧化酶活性,显著降低了叶片内抗氧化酶活性;此外,复合微生物肥料的添加使番茄叶片和根系内MAD含量和O2.-产生速率均显著低于盐胁迫处理,使但是提高了叶片和根系内可溶性蛋白含量增加。以上结果表明,复合微生物肥料可通过调节番茄叶片渗透势、提高根系活力和、光合色素含量以及番茄抗氧化损伤的能力来增强番茄幼苗的抗盐性,通过促进光合制造、贮存碳水化合物的能力来,最终提高番茄幼苗生物量的累积。     

钙和NO对NaCl胁迫下黄瓜幼苗生长和活性氧代谢的影响

采用营养液培养的方法,研究了Ca2+对外源一氧化氮(Nitric oxide,NO)所诱导的NaCl胁迫下黄瓜幼苗生长、活性氧代谢的影响。结果表明,添加外源NO或Ca2+显著缓解了NaCl胁迫对黄瓜幼苗生长的抑制,叶片和根系超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性较单独NaCl胁迫处理显著提高,丙二醛(MDA)和过氧化氢(H2O2)的含量、超氧阴离子(O.-2)产生速率明显下降;添加NO的同时添加Ca2+通道抑制剂La3+抑制了NO的这些调节作用。结果表明Ca2+对NO诱导的NaCl胁迫下黄瓜幼苗植株活性氧清除能力的提高起重要作用,NO的作用可能依赖于胞浆Ca2+。     

NaCl胁迫对菜用大豆种子抗坏血酸-谷胱甘肽循环的影响

采用蛭石栽培,在100 mmol/L NaCl胁迫下,对耐盐性不同的两个菜用大豆[Glycine max(L.)Merr.]品种种子的过氧化氢(H2O2)含量及抗坏血酸-谷胱甘肽(AsA-GSH)循环进行了研究。结果显示,NaCl胁迫显著增加了菜用大豆种子的H2O2含量,但耐盐品种"绿领特早"的增幅低于盐敏感品种"理想高产95-1"。NaCl胁迫期间,"绿领特早"种子中抗坏血酸过氧化物酶(APX)、脱氢抗坏血酸还原酶(DHAR)、谷胱甘肽还原酶(GR)活性,AsA、GSH含量以及AsA/DHA值和GSH/GSSG值的增幅高于同期的"理想高产95-1",或降幅低于同期的"理想高产95-1";脱氢抗坏血酸(DHA)、氧化型谷胱甘肽(GSSG)含量的增幅低于同期的"理想高产95-1"。表明"绿领特早"种子在胁迫期间能够保持较高的AsA-GSH循环效率,可有效地抑制H2O2的积累,这可能是其耐盐性较强的重要原因之一。     

2,4-表油菜素内酯对NaCl胁迫下紫花苜蓿幼苗根系生长抑制及氧化损伤的缓解效应

为探讨外源2,4-表油菜素内酯(2,4-epibrassinolide,EBR)提高紫花苜蓿(Medicago sativa L.)耐盐性的生理调控机制,采用营养液水培法,以紫花苜蓿品种‘中苜3号’和‘陇中苜蓿’为材料,研究Na Cl胁迫下添加外源EBR对紫花苜蓿幼苗根系生长、渗透调节物质含量、抗氧化酶活性、非酶抗氧化物质、活性氧及MDA含量的影响。结果表明:150 mmol·L?1 Na Cl胁迫下,‘中苜3号’、‘陇中苜蓿’幼苗根系的生长显著受到抑制,根系干重和根系活力显著降低,幼苗根系中的可溶性蛋白含量和抗氧化酶活性显著降低,活性氧和MDA含量显著增加,根系脂质过氧化程度加剧,细胞质膜完整性受到破坏。添加0.1μmol·L?1外源EBR显著增加了2个品种苜蓿幼苗的主根长、根系总长度、根系总表面积、根体积、根尖数、根系干重和根系活力,幼苗根系中的可溶性蛋白含量、抗氧化酶(SOD、APX、GPX、CAT、GR)活性、非酶抗氧化物质(As A、GSH)含量均显著增加,超氧阴离子自由基(O2·?)产生速率、羟自由基(OH·)浓度、H2O2含量和膜脂过氧化产物MDA含量显著降低,有效缓解了根系脂质过氧化程度,提高了细胞质膜的完整性。说明外源EBR能够有效缓解Na Cl胁迫对紫花苜蓿幼苗根系生长的抑制作用,增强紫花苜蓿幼苗根系的渗透调节能力和抗氧化系统活性,降低活性氧的积累和膜脂过氧化水平,减小Na Cl胁迫对紫花苜蓿幼苗根系造成的氧化伤害,促进根系生长,增强苜蓿幼苗的抗盐性。     

外源多胺对低氧胁迫下黄瓜幼苗根系活性氧及保护酶活性变化的影响

用亚精胺(Spd)处理了低氧胁迫下的黄瓜幼苗,并对根系中的活性氧及保护酶进行了检测。结果表明,低氧处理后黄瓜幼苗根系的脂质过氧化产物丙二醛(M DA)含量显著升高,活性氧生成速率含量和保护酶活性,都有一个先升后降的过程;Spd处理后,低氧胁迫下的黄瓜幼苗根系中的M DA、活性氧含量显著下降,过氧化物歧化酶(SOD)和过氧化物酶(POD)都保持较高的活性,而过氧化氢酶(CAT)活性较弱。超氧阴离子(O2-)生成速率的变化与SOD活性的变化有一定的相关性,H2O2的含量变化与过氧化物酶(POD)有一定的相关性。     

Physiological and growth responses of two different salt-sensitive cucumber cultivars to NaCl stress

Abstract

The effect of NaCl stress on the growth, membrane permeability, anti-oxidant enzyme activities and ion content of cucumber seedlings was investigated. Two cultivars (Jinchun No. 2, a relatively salt-sensitive cultivar, and Zaoduojia, a relatively salt-tolerant cultivar) of cucumber were used. Shoot and root dry weights, plant height, stem diameter, leaf area and leaf number of both cultivars decreased when NaCl concentrations increased. The decreases in shoot and root dry weights and leaf area were more significant in Jinchun No. 2 than in Zaoduojia. Meanwhile, the salt injury index, the membrane permeability, malondialdehyde (MDA) contents, superoxide dismutase (SOD) and peroxidase (POD) activities of both cultivars increased significantly with salt stress, and the increases in the salt injury index and MDA were higher in Jinchun No. 2 than in Zaoduojia, whereas the increase in POD activity was lower in Jinchun No. 2 than in Zaoduojia. Free proline content of Zaoduojia increased markedly with increasing concentrations of NaCl, whereas the content of Jinchun No. 2 was unaffected by salt stress. In addition, the contents of Na⁺ in the leaf, stem and root of both cultivars increased significantly, whereas the contents of K⁺ decreased significantly, resulting in an increase in the Na⁺/K⁺ ratio when NaCl concentrations increased. These results suggest that Zaoduojia exhibits a better protection mechanism against oxidative damage and lipid peroxidation by maintaining higher proline content and POD activity than the salt-sensitive Jinchun No. 2 cultivar.     

Kinetics of Zinc Uptake and Anatomy of Roots and Leaves of Coffee Trees as Affected by Zinc Nutrition

Abstract

The influence of silicon (Si) (2.5 mM), sodium chloride (NaCl) (100 mM), and Si (2.5 mM) + NaCl (97.5 mM) supply on chlorophyll content, chlorophyll fluorescence, the concentration of malondialdehyde (MDA), H₂O₂ level, and activities of superoxide dismutase (SOD; E.C.1.15.1.1.), ascorbate peroxidase (APx; E.C.1.11.1.11.), catalase (CAT; E.C.1.11.1.6.), guaiacol peroxidase (G-POD; E.C.1.11.1.7.) enzymes, and protein content were studied in tomato (Lycopersicon esculentum Mill c.v.) leaves over 10-day and 27-day periods. The results indicated that silicon partially offset the negative impacts of NaCl stress with increased the tolerance of tomato plants to NaCl salinity by raising SOD and CAT activities, chlorophyll content, and photochemical efficiency of PSII. Salt stress decreased SOD and CAT activities and soluble protein content in the leaves. However, addition of silicon to the nutrient solution enhanced SOD and CAT activities and protein content in tomato leaves under salt stress. In contrast, salt stress slightly promoted APx activity and considerably increased H₂O₂ level and MDA concentration and Si addition slightly decreased APx activity and significantly reduced H₂O₂ level and MDA concentration in the leaves of salt-treated plants. G-POD activity was slightly decreased by addition of salt and Si. Enhanced activities of SOD and CAT by Si addition may protect the plant tissues from oxidative damage induced by salt, thus mitigating salt toxicity and improving the growth of tomato plants. These results confirm that the scavenging system forms the primary defense line in protecting oxidative damage under stress in crop plants.     

硅促进盐胁迫下黄瓜NHX1基因表达及Na+在液泡中的区隔化效应

  【目的】   硅可提高植物的耐盐性,但不同植物中硅提高耐盐性的机理并不相同。探究硅对盐胁迫下黄瓜幼苗的氧化损伤、Na+积累和激素水平的影响,以阐明硅提高黄瓜耐盐性的机制。   【方法】   以基因型为Mch-4的黄瓜幼苗为试材,进行水培试验。营养液中NaCl的胁迫浓度为65 mmol/L,施硅水平为Na₂SiO₃·9H₂O 0.3 mmol/L。在处理10天后,测定黄瓜幼苗生物量、Na+含量与分配、Na+转运相关基因表达水平及激素含量。   【结果】   施硅可改善盐胁迫下黄瓜幼苗的生长,减轻植株的氧化损伤。硅对盐胁迫下黄瓜根系和叶片Na+含量无明显影响,可显著降低根和叶中质膜Na+/H+反向转运蛋白基因SOS1的表达量,对高亲和力钾转运蛋白基因HKT1的表达均影响不大,但促进了液泡膜Na+/H+反向转运蛋白基因NHX1的表达。对盐胁迫下黄瓜叶片Na+的亚细胞定位发现,硅处理使叶绿体中Na+含量下降,而液泡中Na+含量升高。硅处理提高了盐胁迫植株根和叶片中赤霉素、生长素和细胞分裂素的水平。   【结论】   施硅可提高液泡膜Na+/H+反向转运蛋白基因NHX1的表达,将Na+区隔化于液泡中,进而降低叶绿体中的Na+含量,缓解盐胁迫下黄瓜幼苗的氧化损伤;硅还诱导产生了较多的赤霉素、生长素和细胞分裂素,其调控Na+积累和黄瓜幼苗的氧化损伤的机理还需进一步研究。     

Protective role of proline against salt stress is partially related to the improvement of water status and peroxidase enzyme activity in cucumber

A salt-sensitive cucumber cultivar "Jinchun No. 2" ( Cucumis sativus L.) was used to investigate the role of proline in alleviating salt stress in cucumber. Proline was applied twice (day 0 and day 4 after salt treatment) as a foliar spray, with a volume of 25 mL per plant at each time. Plant dry weight, leaf relative water content, proline, malondialdehyde (MDA), Na⁺, K⁺ and Cl⁻ contents, as well as superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) activities in the plants were determined at day 8 after salt treatment. The results showed that 100 mmol L^–1 NaCl stress significantly decreased plant dry weight, leaf relative water and K⁺ contents, and increased leaf MDA, Na⁺ and Cl⁻ contents and SOD, POD, CAT and APX activities. However, leaf proline accumulation was not affected by salinity. The exogenous application of proline significantly alleviated the growth inhibition of plants induced by NaCl, and was accompanied by higher leaf relative water content and POD activity, higher proline and Cl⁻ contents, and lower MDA content and SOD activity. However, there was no significant difference in Na⁺ and K⁺ contents or in CAT and APX activities between proline-treated and untreated plants under salt stress. Taken together, these results suggested that the foliar application of proline was an effective way to improve the salt tolerance of cucumber. The enhanced salt tolerance could be partially attributed to the improved water status and peroxidase enzyme activity in the leaf.     

H+‐atpase and H+‐transport activities in tonoplast vesicles from barley roots under salt stress and influence of calcium and abscisic acid 1

Seedlings of two barley cultivars differing in NaCl sensitivity were treated with low (100 mM) or high (400 mM) concentration of NaCl for 6 days. Tonoplast vesicles were prepared from roots, and H⁺‐ATPase and H⁺‐transport activities associated with tonoplast were assayed. Both H⁺‐ATPase and H⁺‐transport activities in the two cultivars were increased at 100 mM NaCl. These activities also increased in the salt‐tolerant cultivar at 400 mM NaCl, but in salt‐sensitive cultivar were decreased. In vivo treatment with 10 mM Ca²⁺ stimulated H⁺‐ATPase and H⁺‐transport activities at two levels of NaCl, however, treatment with 10⁵M (±) abscisic acid (ABA) inhibited these activities. From these results we propose that the increase of the vacuolar H⁺ pumps in barley roots reflects an adaptation to salt stress. The stimulation of HVATPase and H⁺‐transport activities by calcium (Ca) depends mainly on its effect in maintaining stability of membrane under salt stress.     

低钾对水稻不同叶位叶片光合特性及抗氧化系统的影响

研究了低钾条件下钾敏感型水稻品种"二九丰"不同叶位叶片的气体交换、荧光参数、色素和活性氧及抗氧化酶活性的变化。结果表明,随着叶位降低,对照和低钾处理的净光合速率(Pn)、气孔导度(Gs)、叶绿素(Chl)、类胡萝卜素(Car)、PSⅡ最大光化学量子产量(Fv/Fm)、PSⅡ实际光化学量子产量(ΦPSⅡ)、表观电子传递速率(ETR)、可溶性蛋白(SP)含量、超氧化物歧化酶(SOD)和过氧化物酶(POD)活性均显著下降,而胞间CO2浓度(Ci)、激发压(1-qP)、非光化学猝灭(qN)、超氧化物自由基(O2.-)产生速率、过氧化氢(H2O2)和丙二醛(MDA)含量却急剧升高。与对照相比,低钾胁迫加速了各叶位Pn的下降和Ci的上升,而Gs并无降低,说明Pn的降低主要是由非气孔引起的。低钾胁迫还明显降低Fv/Fm、ΦPSⅡ、ETR、Chl和SP含量、Chl/Car比率、及SOD和POD活性,促进了(1-qP)、O2.-产生速率、H2O2和MDA含量的升高。这些结果均说明低钾胁迫引起了PSⅡ光化学效率降低和能量耗散的不平衡,从而导致了叶片的早衰。相关性分析表明,PSⅡ的激发压的升高与Pn、Chl和SP的含量及SOD和POD活性呈极显著负相关,与O2.-产生速率、H2O2和MDA含量呈极显著的正相关。据此推测,激发压的升高可能介导了水稻在低钾条件下叶片早衰的起始过程。     

Protective role of proline against salt stress is partially related to the improvement of water status and peroxidase enzyme activity in cucumber

Abstract

A salt-sensitive cucumber cultivar “Jinchun No. 2” (Cucumis sativus L.) was used to investigate the role of proline in alleviating salt stress in cucumber. Proline was applied twice (day 0 and day 4 after salt treatment) as a foliar spray, with a volume of 25?mL per plant at each time. Plant dry weight, leaf relative water content, proline, malondialdehyde (MDA), Na⁺, K⁺ and Cl^? contents, as well as superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) activities in the plants were determined at day 8 after salt treatment. The results showed that 100?mmol?L^–1 NaCl stress significantly decreased plant dry weight, leaf relative water and K⁺ contents, and increased leaf MDA, Na⁺ and Cl^? contents and SOD, POD, CAT and APX activities. However, leaf proline accumulation was not affected by salinity. The exogenous application of proline significantly alleviated the growth inhibition of plants induced by NaCl, and was accompanied by higher leaf relative water content and POD activity, higher proline and Cl^? contents, and lower MDA content and SOD activity. However, there was no significant difference in Na⁺ and K⁺ contents or in CAT and APX activities between proline-treated and untreated plants under salt stress. Taken together, these results suggested that the foliar application of proline was an effective way to improve the salt tolerance of cucumber. The enhanced salt tolerance could be partially attributed to the improved water status and peroxidase enzyme activity in the leaf.     

盐胁迫下大气CO2浓度升高对黄瓜幼苗生长、光合特性及矿质养分吸收的影响

采用营养液培养和开顶箱法,研究了盐胁迫下CO2浓度升高对黄瓜幼苗生长、光合特性及矿质养分吸收的影响。结果表明,黄瓜生长在80mmol/L NaCl下,其生物量、光合速率、气孔导度、蒸腾速率均显著下降,而胞间CO2浓度明显升高;CO2浓度升高可增加盐胁迫下黄瓜幼苗生物量,使光合速率、气孔导度、蒸腾速率升高。表明CO2浓度升高能减轻盐胁迫对光合功能的不利效应。80mmol/L NaCl可使黄瓜幼苗体内总N和K 的浓度降低,而使Na 浓度增加;CO2浓度升高具有提高盐胁迫下总N和K 浓度,降低Na 浓度的效应,说明CO2浓度升高可减轻盐胁迫的毒害作用,提高黄瓜幼苗生物量。