绵羊口液对羊草生长的作用机制研究
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摘要
动植物关系（plant-animal relationship）是生态学研究的核心问题之一。在自然界长期的协同进化过程中，动植物之间形成了稳定的适应（adaptation）和互惠关系（mutualism）。动物采食对植物的影响包括机械损伤（mechanical damage）和口液作用（saliva effect）。动物口液中的信号物质可以诱导植物化学防御（chemical defense），同时也会调节植物的生理代谢过程，促进植物生长。动物口液对植物生长的刺激作用是植物对动物采食补偿性响应的重要机制。因此，研究动物口液对植物生长的影响及其作用机制，对于理解动植物之间的协同进化和互惠关系具有重要意义。目前关于动物口液对植物生长影响的研究较少，机制尚不清楚。
据此，本论文以松嫩草原优势物种（羊草，Leymus chinensis (Trin.) Tzvelev.）和主要放牧家畜（绵羊）为研究对象，分析探讨了动物口液对植物生长的作用规律及内在机制，具体包括以下5方面的研究内容：（1）动物口液是否能够促进植物生长？（2）动物采食强度对口液作用效果的影响；（3）口液作用随时间的变化规律；（4）口液中的重要化学成分对植物生长的影响；（5）植物碳水化合物转运对动物口液的响应。
本研究通过盆栽的控制实验，采用完全随机实验设计和高效液相色谱分析技术，对绵羊口液调节羊草生长的作用规律及机制进行深入探讨，获得了以下主要结论：
动物口液能够促进植物生长，并且口液对植物生长的促进作用主要表现为能够增加分蘖和芽库的数量。三年（2006、2007和2008）的实验结果表明，口液能够增加羊草地上生物量、地下生物量、分蘖和芽库数量，而对羊草高度没有显著作用。口液对羊草生长的促进作用主要表现在提高其分蘖能力上。羊草等多年生禾本科植物的生长过程包括伸长生长和增加分蘖两个阶段，伸长生长和分蘖这两种生长方式在时间上存在差异。植物生长发育早期进行伸长生长，生长达到一定阶段后，伸长生长逐渐减慢甚至停止，地下根茎上的芽库开始输出，不断增加地上分蘖。本研究进行实验处理时，羊草已开始分蘖，而伸长生长基本停止，所以芽库数量和分蘖对口液作用的响应最为敏感。分蘖是羊草的主要繁殖方式，也是对动物采食的重要适应性特征，芽库数量决定羊草潜在的分蘖能力。因此，口液对羊草分蘖和芽库的促进作用，能够提高羊草对动物采食的耐受性，维持动植物之间稳定的共存关系。
动物口液对植物生长的作用具有采食强度依赖性，在轻度和完全刈割水平上，口液的作用最强。通过比较不同刈割水平上（0%，25%，50%，75%和100%）绵羊口液对羊草生长的作用差异，发现在轻度刈割（25%）和完全刈割（100%）两个刈割水平上，口液对羊草生长的促进作用最强。这是因为两个刈割水平分别接近羊草的顶端分生组织和基部分生组织，而口液中的生长因子和生长调节物质在分生组织附近对植物生长的调节作用最为有效。多年生禾本科植物的生长发育受顶端分生组织、居间分生组织和基部分生组织的调节，而植物的分蘖动态由顶端分生组织和基部分生组织的控制。本研究中，完全刈割水平（100%）靠近地面，与基部分生组织的距离最近，口液中的生长调节物质最容易到达地下芽库，能够刺激芽库输出，进而促进羊草进行分蘖。轻度刈割水平（25%）接近顶端分生组织，口液中的生长因子会通过顶端分生组织调节植物分蘖动态。因此，绵羊口液对羊草生长的作用具有刈割强度依赖性，在轻度和完全两个刈割水平上作用最强。在轻度采食强度上，动物口液可以提高植物的补偿生长能力，为优化放牧假说（optimal grazing hypothesis）提供新的实验支持证据；而在100%的完全采食强度上，口液对植物生长的促进作用可以降低重度放牧对植物的损伤，提高植物对动物采食的耐受性及其对长期放牧的适应性。
动物口液对植物生长的调节作用具有时间依赖性。在本研究中，处理半天内羊草各构件内碳水化合物便发生变化；处理10天后口液对植株高度的影响已经表现出来；而口液对羊草分蘖和芽库数量以及生物量的作用主要表现在处理一个月后，其后，口液作用逐渐消失。植物被动物采食后，其再生能力受到贮藏碳水化合物转运和分生组织活性的调节。碳水化合物转运等生理过程在短时间内就会对动物采食做出响应，而分生组织对植物生长的调节过程需要较长的时间，所以口液对植物生长特征的影响需要更长的作用时间。植物伸长生长和分蘖两种生长方式对动物采食的敏感程度不同，伸长生长较快，而休眠芽库的激活需要一定诱导时间（inductive timeperiod）。因此，动物口液对植物生长的促进作用具有时间依赖性（time dependence），并且口液对植物生理过程和生长特征的作用时间存在差异。口液作用的时间依赖性表明，植物被动物采食后需要一定的时间进行恢复生长，这一研究结论强调轮牧、休牧等放牧管理策略的重要性。
口液成分对植物生长没有发现促进作用。本研究比较了绵羊口液，表皮生长因子（epidermal growth factor）和硫胺素（thiamine），以及两者混合物对羊草生长的影响。实验结果表明，表皮生长因子和硫胺素，以及两者的混合物都没有促进羊草生长，而绵羊口液对羊草生长具有促进作用。表皮生长因子和硫胺素对羊草生长没有促进作用，可能是由于羊草生长在营养丰富的环境中，其合成的生长调节物质足以满足自身需要，而施加的外源生长物质对植物生长未能发挥促进作用。
动物口液可以促进植物碳水化合物的转运。贮藏碳水化合物的快速转运是植物耐受动物采食、提高补偿生长能力的重要机制。本研究的实验结果显示，刈割处理3天后羊草各构件内果聚糖含量降低30-50%，之后趋于稳定。叶片和茎中的蔗糖含量在处理1天后增加2-3倍，3天后其含量降低到处理前的水平，而根茎和须根中的蔗糖含量在处理之后逐渐升高到处理前的3倍左右；刈割后，羊草各构件内葡糖和果糖含量逐渐增加到处理前的2-10倍。相对于单独刈割，绵羊口液加快了羊草各构件果聚糖的分解和单糖的积累。结果表明，被动物采食后的短时间内，植物各构件内的聚糖逐渐分解，单糖大量积累，为植物新生组织的再生过程提供能量。动物对植物碳水化合物转运的促进作用是植物补偿性生长的重要生理机制。
总之，通过系列控制实验，本研究对动物口液影响植物生长的作用规律及机制获得了新的认识：绵羊口液能够促进羊草生长，并且口液作用具有采食强度和时间依赖性；采食强度对口液作用的影响是由植物分生组织的分布规律决定的，在顶端分生组织和基分生组织的位置上，动物口液对植物生长的调节作用最为有效。在轻度采食强度上动物口液可以提高植物的超补偿生长能力，这为优化放牧假说提供新的实验证据；在完全采食强度上，口液对植物生长的促进作用可以提高植物对重度放牧的适应性。动物口液对植物生长调节作用具有时间依赖性，这表明植物被动物采食后需要一定的恢复时间，这一结论强调休牧、轮牧等放牧管理策略的重要性。动物口液能够加快植物被动物采食后再生过程中碳水化合物的转运，为植物再生生长提供能量，这是植物对动物采食补偿生长的重要生理机制。本研究阐明了动物口液对植物生长的作用规律及机制，加深了对植物采食耐受性和补偿性生长机制的理解，丰富了动植物之间的互惠和协同进化理论，并为制定合理的草地管理策略提供科学依据。
Plant-animal relationship is a critical issue in ecology, and with long co-evolutionaryhistory there is complex adaptation and mutualism between grazer and grass in grasslandecosystems. Impacts of foraging on plant include influence of mechanical damage andsaliva effects. Cues in animal saliva can induce plant chemical defense, and regulatephysiological process of plant regrowth. The stimulatory effect of animal saliva is one ofthe mechanisms for plant compensatory response to herbivory. Therefore, the study onsalivary effects on plant growth and the mechanism can improve our knowledge on theco-evolutionary and mutualistic association between plants and animals. However, thereis little work on the impact of saliva on plant growth, and the mechanism keeps unclear.
We performed a set of experiments to explore the effects of sheep saliva on growthof Leymus chinensis (Trin.) Tzvelev., which is a dominate species in the SongnenGrassland. This study was conducted to examine,(1) impacts of animal saliva on plantgrowth,(2) influence of herbivory intensity on saliva impacts,(3) change of saliva effectswith time,(4) response of plant growth to chemical components of animal saliva, and (5)effects of animal saliva on remobilization of carbohydrate reserve.
Experiments were conducted in controlled environments with completelyrandomized design and carbohydrate was measured with high performance liquidchromatography (HPLC). The main results and conclusions are present as following:
Sheep saliva stimulated growth of L. chinensis, especially for tillers and buds.Results in2006and2007showed that sheep saliva promoted growth of L. chinensis,increasing biomass, buds and tillers, but had no influence on height. Recruitment ofperennial growth depends mainly on vegetative growth, including elongation and tillering.Elongation is regulated by intercalary meristem and dynamic of tillers results fromoutgrowth of buds. The two growth patterns work at different time within one growingseason. At early period of development, plant growth comes from shoot elongation, inwhich there is accumulative increase in plant height and leaf area, and consequent highcapacity of photosynthesis and accumulation of assimilate. In the middle of development,elongation declines and even stops, and belowground buds begin to outgrowth. Whentreatments were performed in this study, L. chinensis began to tiller, and at this time basalmeristem started to regulate plant growth. In results of this study, tillers and budsresponded sensitively to saliva application, which can enhance plant tolerance toherbivory and keep the coexistence between plant and animal.
Impact of animal saliva on plant growth was herbivory intensity dependent, andat light and complete herbivory intensities it was strongest. Results in three yearsshowed that sheep saliva had higher impacts on growth of L. chinensis at light (25%) andcomplete clipping (100%) levels. We suggested that this was associated with meristem distribution pattern within perennial herbaceous plants. Development of perennial grasseswas regulated by apical meristem, intercalary meristem and basal meristem. Intercalarymeristem controls plant elongation, and basal meristem and rhizomatous buds determinedynamic of tillering. Activity of both the meristem is regulated by apical meristem. Theresults showed that at light and complete clipping levels, sheep saliva had strongestpromotive effects on plant growth because the two positions were most close to apical andbasal meristem respectively. Tillers of L. chinensis came from outgrowth of buds at thebase of shoot and along the rhizome. The complete clipping level was most close to theground, and so it was easiest for growth regulators in animal saliva to reach belowgroundbud bank and stimulate tillering. Light clipping level was close to apical meristem andanimal saliva could regulate tillering through the regulation on apical meristem. Therefore,effect of sheep saliva on growth of L. chinensis was clipping level dependent, and at lightand complete clipping levels it was strongest.
Effect of animal saliva on plant growth was time dependent. In this study, therewas response in carbohydrate content to sheep saliva within half a day after treatments.Plant height responded to saliva10days after treatments and30days after treatments,there was significant response in tillers, buds and biomass. Regrowth capacity of plantfollowing herbivory depends on the remobilization of restored carbohydrate and activityof meristem. Transfer of carbon reserve can respond to herbivory in a short time, andplant elongation and tillering differs in the sensitivity to herbivory. In this study,carbohydrate in every plant part changed within half a day after treatments, ten days laterthere was response to plant height, and thirty days after treatments impacts of sheep salivaon tillers and buds emerged. So, impact of animal saliva on plant growth was timedependent and there was difference in plant growth parameters to respond to saliva.
The main salivary components had no positive effects on plant growth. There isa huge amount of growth factors and regulators within animal saliva, and saliva plays animportant role in plant-animal interaction. In this study, we compared different responseof L. chinensis growth to sheep saliva, epidermal growth factor (EGF), thiamine and themixture of the two chemicals. Results showed that EGF, thiamine or their mixture did notpromote growth of L. chinensis, and sheep saliva had high promotive effects than EGF,thiamine and their mixture. EGF or thiamine had no stimulation on growth of L. chinensis,and this may result from the favourable growth condition, in which plant growth is notlimited by growth factors.
Animal saliva promoted remobilization of carbohydrate reserve. Rapidremobilization of carbohydrate reserve is one of the mechanisms of plant tolerance toherbivory and its capacity of compensatory growth. Results of this study showed that inevery plant part fructan content decreased by30-50%three days after treatments, and thekept in the later period of the experiment. Sucrose in leaf and stem increased by2to3times one days after treatments and then decreased to the level before treatments, while inrhizome and fine root sucrose content increased gradually three times. In every plant part,concentration of glucose and fructose increased gradually by2to10times. Compared with clipping, sheep saliva promoted hydrolization of fructans and accumulation ofglucose and fructose in every plant part, while it suppressed the accumulation of sucroseand then improved its concentration. The results demonstrated that during the short periodafter herbivory, fructans decreased, monosaccharides increased and were transported intonew tissues. In the process of herbivory, the saliva deposited on plant surface promotedmobilization of carbohydrate to support energy for compensatory growth.
Results of this study showed that sheep saliva promoted growth and saliva effectswere herbivory intensity dependent. We proposed that this was associated withdistribution pattern of meristem within plant, and at the point of apical and basal meristemthe effects of animal saliva were most effective. The promotive effect of saliva on plantgrowth at intensive herbivory levels was associated with plant adaptation to long termgrazing, and the stimulatory impact of saliva on plant compensation at light herbivoryintensity can increase plant compensatory response to herbivory that gave support to theoptimal grazing hypothesis. The time dependence of animal saliva on plant growthaddressed the significance of delaying grazing and rotational grazing. Animal salivapromoted remobilization of carbohydrate during regrowth, with provided physiologicalmechanism of plant compensatory response and tolerance to herbivory. The stimulatoryeffects of animal saliva on plant growth and remobilization of carbohydrate expanded themutualism and coevolution theory, and gave guide to scientific grassland management.
据此，本论文以松嫩草原优势物种（羊草，Leymus chinensis (Trin.) Tzvelev.）和主要放牧家畜（绵羊）为研究对象，分析探讨了动物口液对植物生长的作用规律及内在机制，具体包括以下5方面的研究内容：（1）动物口液是否能够促进植物生长？（2）动物采食强度对口液作用效果的影响；（3）口液作用随时间的变化规律；（4）口液中的重要化学成分对植物生长的影响；（5）植物碳水化合物转运对动物口液的响应。
本研究通过盆栽的控制实验，采用完全随机实验设计和高效液相色谱分析技术，对绵羊口液调节羊草生长的作用规律及机制进行深入探讨，获得了以下主要结论：
动物口液能够促进植物生长，并且口液对植物生长的促进作用主要表现为能够增加分蘖和芽库的数量。三年（2006、2007和2008）的实验结果表明，口液能够增加羊草地上生物量、地下生物量、分蘖和芽库数量，而对羊草高度没有显著作用。口液对羊草生长的促进作用主要表现在提高其分蘖能力上。羊草等多年生禾本科植物的生长过程包括伸长生长和增加分蘖两个阶段，伸长生长和分蘖这两种生长方式在时间上存在差异。植物生长发育早期进行伸长生长，生长达到一定阶段后，伸长生长逐渐减慢甚至停止，地下根茎上的芽库开始输出，不断增加地上分蘖。本研究进行实验处理时，羊草已开始分蘖，而伸长生长基本停止，所以芽库数量和分蘖对口液作用的响应最为敏感。分蘖是羊草的主要繁殖方式，也是对动物采食的重要适应性特征，芽库数量决定羊草潜在的分蘖能力。因此，口液对羊草分蘖和芽库的促进作用，能够提高羊草对动物采食的耐受性，维持动植物之间稳定的共存关系。
动物口液对植物生长的作用具有采食强度依赖性，在轻度和完全刈割水平上，口液的作用最强。通过比较不同刈割水平上（0%，25%，50%，75%和100%）绵羊口液对羊草生长的作用差异，发现在轻度刈割（25%）和完全刈割（100%）两个刈割水平上，口液对羊草生长的促进作用最强。这是因为两个刈割水平分别接近羊草的顶端分生组织和基部分生组织，而口液中的生长因子和生长调节物质在分生组织附近对植物生长的调节作用最为有效。多年生禾本科植物的生长发育受顶端分生组织、居间分生组织和基部分生组织的调节，而植物的分蘖动态由顶端分生组织和基部分生组织的控制。本研究中，完全刈割水平（100%）靠近地面，与基部分生组织的距离最近，口液中的生长调节物质最容易到达地下芽库，能够刺激芽库输出，进而促进羊草进行分蘖。轻度刈割水平（25%）接近顶端分生组织，口液中的生长因子会通过顶端分生组织调节植物分蘖动态。因此，绵羊口液对羊草生长的作用具有刈割强度依赖性，在轻度和完全两个刈割水平上作用最强。在轻度采食强度上，动物口液可以提高植物的补偿生长能力，为优化放牧假说（optimal grazing hypothesis）提供新的实验支持证据；而在100%的完全采食强度上，口液对植物生长的促进作用可以降低重度放牧对植物的损伤，提高植物对动物采食的耐受性及其对长期放牧的适应性。
动物口液对植物生长的调节作用具有时间依赖性。在本研究中，处理半天内羊草各构件内碳水化合物便发生变化；处理10天后口液对植株高度的影响已经表现出来；而口液对羊草分蘖和芽库数量以及生物量的作用主要表现在处理一个月后，其后，口液作用逐渐消失。植物被动物采食后，其再生能力受到贮藏碳水化合物转运和分生组织活性的调节。碳水化合物转运等生理过程在短时间内就会对动物采食做出响应，而分生组织对植物生长的调节过程需要较长的时间，所以口液对植物生长特征的影响需要更长的作用时间。植物伸长生长和分蘖两种生长方式对动物采食的敏感程度不同，伸长生长较快，而休眠芽库的激活需要一定诱导时间（inductive timeperiod）。因此，动物口液对植物生长的促进作用具有时间依赖性（time dependence），并且口液对植物生理过程和生长特征的作用时间存在差异。口液作用的时间依赖性表明，植物被动物采食后需要一定的时间进行恢复生长，这一研究结论强调轮牧、休牧等放牧管理策略的重要性。
口液成分对植物生长没有发现促进作用。本研究比较了绵羊口液，表皮生长因子（epidermal growth factor）和硫胺素（thiamine），以及两者混合物对羊草生长的影响。实验结果表明，表皮生长因子和硫胺素，以及两者的混合物都没有促进羊草生长，而绵羊口液对羊草生长具有促进作用。表皮生长因子和硫胺素对羊草生长没有促进作用，可能是由于羊草生长在营养丰富的环境中，其合成的生长调节物质足以满足自身需要，而施加的外源生长物质对植物生长未能发挥促进作用。
动物口液可以促进植物碳水化合物的转运。贮藏碳水化合物的快速转运是植物耐受动物采食、提高补偿生长能力的重要机制。本研究的实验结果显示，刈割处理3天后羊草各构件内果聚糖含量降低30-50%，之后趋于稳定。叶片和茎中的蔗糖含量在处理1天后增加2-3倍，3天后其含量降低到处理前的水平，而根茎和须根中的蔗糖含量在处理之后逐渐升高到处理前的3倍左右；刈割后，羊草各构件内葡糖和果糖含量逐渐增加到处理前的2-10倍。相对于单独刈割，绵羊口液加快了羊草各构件果聚糖的分解和单糖的积累。结果表明，被动物采食后的短时间内，植物各构件内的聚糖逐渐分解，单糖大量积累，为植物新生组织的再生过程提供能量。动物对植物碳水化合物转运的促进作用是植物补偿性生长的重要生理机制。
总之，通过系列控制实验，本研究对动物口液影响植物生长的作用规律及机制获得了新的认识：绵羊口液能够促进羊草生长，并且口液作用具有采食强度和时间依赖性；采食强度对口液作用的影响是由植物分生组织的分布规律决定的，在顶端分生组织和基分生组织的位置上，动物口液对植物生长的调节作用最为有效。在轻度采食强度上动物口液可以提高植物的超补偿生长能力，这为优化放牧假说提供新的实验证据；在完全采食强度上，口液对植物生长的促进作用可以提高植物对重度放牧的适应性。动物口液对植物生长调节作用具有时间依赖性，这表明植物被动物采食后需要一定的恢复时间，这一结论强调休牧、轮牧等放牧管理策略的重要性。动物口液能够加快植物被动物采食后再生过程中碳水化合物的转运，为植物再生生长提供能量，这是植物对动物采食补偿生长的重要生理机制。本研究阐明了动物口液对植物生长的作用规律及机制，加深了对植物采食耐受性和补偿性生长机制的理解，丰富了动植物之间的互惠和协同进化理论，并为制定合理的草地管理策略提供科学依据。
Plant-animal relationship is a critical issue in ecology, and with long co-evolutionaryhistory there is complex adaptation and mutualism between grazer and grass in grasslandecosystems. Impacts of foraging on plant include influence of mechanical damage andsaliva effects. Cues in animal saliva can induce plant chemical defense, and regulatephysiological process of plant regrowth. The stimulatory effect of animal saliva is one ofthe mechanisms for plant compensatory response to herbivory. Therefore, the study onsalivary effects on plant growth and the mechanism can improve our knowledge on theco-evolutionary and mutualistic association between plants and animals. However, thereis little work on the impact of saliva on plant growth, and the mechanism keeps unclear.
We performed a set of experiments to explore the effects of sheep saliva on growthof Leymus chinensis (Trin.) Tzvelev., which is a dominate species in the SongnenGrassland. This study was conducted to examine,(1) impacts of animal saliva on plantgrowth,(2) influence of herbivory intensity on saliva impacts,(3) change of saliva effectswith time,(4) response of plant growth to chemical components of animal saliva, and (5)effects of animal saliva on remobilization of carbohydrate reserve.
Experiments were conducted in controlled environments with completelyrandomized design and carbohydrate was measured with high performance liquidchromatography (HPLC). The main results and conclusions are present as following:
Sheep saliva stimulated growth of L. chinensis, especially for tillers and buds.Results in2006and2007showed that sheep saliva promoted growth of L. chinensis,increasing biomass, buds and tillers, but had no influence on height. Recruitment ofperennial growth depends mainly on vegetative growth, including elongation and tillering.Elongation is regulated by intercalary meristem and dynamic of tillers results fromoutgrowth of buds. The two growth patterns work at different time within one growingseason. At early period of development, plant growth comes from shoot elongation, inwhich there is accumulative increase in plant height and leaf area, and consequent highcapacity of photosynthesis and accumulation of assimilate. In the middle of development,elongation declines and even stops, and belowground buds begin to outgrowth. Whentreatments were performed in this study, L. chinensis began to tiller, and at this time basalmeristem started to regulate plant growth. In results of this study, tillers and budsresponded sensitively to saliva application, which can enhance plant tolerance toherbivory and keep the coexistence between plant and animal.
Impact of animal saliva on plant growth was herbivory intensity dependent, andat light and complete herbivory intensities it was strongest. Results in three yearsshowed that sheep saliva had higher impacts on growth of L. chinensis at light (25%) andcomplete clipping (100%) levels. We suggested that this was associated with meristem distribution pattern within perennial herbaceous plants. Development of perennial grasseswas regulated by apical meristem, intercalary meristem and basal meristem. Intercalarymeristem controls plant elongation, and basal meristem and rhizomatous buds determinedynamic of tillering. Activity of both the meristem is regulated by apical meristem. Theresults showed that at light and complete clipping levels, sheep saliva had strongestpromotive effects on plant growth because the two positions were most close to apical andbasal meristem respectively. Tillers of L. chinensis came from outgrowth of buds at thebase of shoot and along the rhizome. The complete clipping level was most close to theground, and so it was easiest for growth regulators in animal saliva to reach belowgroundbud bank and stimulate tillering. Light clipping level was close to apical meristem andanimal saliva could regulate tillering through the regulation on apical meristem. Therefore,effect of sheep saliva on growth of L. chinensis was clipping level dependent, and at lightand complete clipping levels it was strongest.
Effect of animal saliva on plant growth was time dependent. In this study, therewas response in carbohydrate content to sheep saliva within half a day after treatments.Plant height responded to saliva10days after treatments and30days after treatments,there was significant response in tillers, buds and biomass. Regrowth capacity of plantfollowing herbivory depends on the remobilization of restored carbohydrate and activityof meristem. Transfer of carbon reserve can respond to herbivory in a short time, andplant elongation and tillering differs in the sensitivity to herbivory. In this study,carbohydrate in every plant part changed within half a day after treatments, ten days laterthere was response to plant height, and thirty days after treatments impacts of sheep salivaon tillers and buds emerged. So, impact of animal saliva on plant growth was timedependent and there was difference in plant growth parameters to respond to saliva.
The main salivary components had no positive effects on plant growth. There isa huge amount of growth factors and regulators within animal saliva, and saliva plays animportant role in plant-animal interaction. In this study, we compared different responseof L. chinensis growth to sheep saliva, epidermal growth factor (EGF), thiamine and themixture of the two chemicals. Results showed that EGF, thiamine or their mixture did notpromote growth of L. chinensis, and sheep saliva had high promotive effects than EGF,thiamine and their mixture. EGF or thiamine had no stimulation on growth of L. chinensis,and this may result from the favourable growth condition, in which plant growth is notlimited by growth factors.
Animal saliva promoted remobilization of carbohydrate reserve. Rapidremobilization of carbohydrate reserve is one of the mechanisms of plant tolerance toherbivory and its capacity of compensatory growth. Results of this study showed that inevery plant part fructan content decreased by30-50%three days after treatments, and thekept in the later period of the experiment. Sucrose in leaf and stem increased by2to3times one days after treatments and then decreased to the level before treatments, while inrhizome and fine root sucrose content increased gradually three times. In every plant part,concentration of glucose and fructose increased gradually by2to10times. Compared with clipping, sheep saliva promoted hydrolization of fructans and accumulation ofglucose and fructose in every plant part, while it suppressed the accumulation of sucroseand then improved its concentration. The results demonstrated that during the short periodafter herbivory, fructans decreased, monosaccharides increased and were transported intonew tissues. In the process of herbivory, the saliva deposited on plant surface promotedmobilization of carbohydrate to support energy for compensatory growth.
Results of this study showed that sheep saliva promoted growth and saliva effectswere herbivory intensity dependent. We proposed that this was associated withdistribution pattern of meristem within plant, and at the point of apical and basal meristemthe effects of animal saliva were most effective. The promotive effect of saliva on plantgrowth at intensive herbivory levels was associated with plant adaptation to long termgrazing, and the stimulatory impact of saliva on plant compensation at light herbivoryintensity can increase plant compensatory response to herbivory that gave support to theoptimal grazing hypothesis. The time dependence of animal saliva on plant growthaddressed the significance of delaying grazing and rotational grazing. Animal salivapromoted remobilization of carbohydrate during regrowth, with provided physiologicalmechanism of plant compensatory response and tolerance to herbivory. The stimulatoryeffects of animal saliva on plant growth and remobilization of carbohydrate expanded themutualism and coevolution theory, and gave guide to scientific grassland management.
引文
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