刈割时间、刈割强度与施肥处理对燕麦补偿的影响

刈割强度、刈割时间和施肥状况对燕麦的补偿有明显影响作用。在不施肥条件下,分蘖期轻度刈割有利于植物的补偿作用,拔节期重度刈割以及重复刈割影响植物生长。施肥可以提高受适度刈割损害植物的补偿程度。无论施肥与否,燕麦都没有发生明显的超补偿现象。但在施肥条件下,留茬高度8cm的刈割处理使燕麦在一定程度上提高了植物的生产力,尤其是秆叶的干重。     

种群密度与施肥对垂穗披碱草刈割后补偿作用的影响

 以甘南亚高山草甸常见牧草垂穗披碱草(Elymus nutans)为对象,通过考察种群密度、施肥与刈割处理等对植物生长和生殖的影响效应,比较了垂穗披碱草在5个密度及2个施肥实验处理条件下对4种刈割处理的补偿性反应特点。结果表明,在所有密度及施肥处理条件下,早期轻度刈割处理 (分蘖期刈割,留茬4 cm) 都有利于植物的补偿作用,后期重度处理 (拔节期刈割,留茬2 cm) 可显著降低植物的地上部分生物量及生殖部分干重 (穗重)。在不施肥情况下,刈割对垂穗披碱草的影响程度随种群密度而加大,在低密度处理中早期轻度刈割的植物发生了超补偿。可以认为,低密度种群中植物具有较多的分蘖是植物在刈割后表现出较高补偿能力的一个重要生物学原因。实验还发现,施肥可提高植物个体的分蘖能力和秆叶再生能力,因而总体上可增强植物的补偿能力,并且在中等密度条件下垂穗披碱草的补偿能力较强,尽管没有发生超补偿现象。不过,施肥主要是有利于植物地上营养器官的补偿,生殖器官的补偿程度较小一些。试验结果对科学管理人工草场具有一定的指导意义。     

施肥和刈割对冷地早熟禾补偿生长的影响

以青藏高原东部高寒草甸常见牧草冷地早熟禾(Poa crymophila)为研究对象,比较研究了肥力和不同刈割处理对植物生长和生物量补偿的影响。实验采用随机设计,肥力分为施肥和不施肥,刈割处理为刈割时间、刈割频度和刈割强度3因子组合。结果表明:不刈割情况下,施肥显著促进冷地早熟禾的生长,而在刈割情况下,施肥对植物的生长没有明显的促进作用甚至有负作用。不施肥,刈割可促进冷地早熟禾地上生物量的增加并发生超补偿,但补偿程度因刈割强度、刈割频度和刈割时间而异;施肥时,刈割没有或很少促进植株发生超补偿。超补偿发生时,资源贫瘠的条件可能是必需的。冷地早熟禾发生超补偿可能是以损耗地下部分的资源为代价的,刈割刺激分蘖的增加对超补偿的发生也有一定的贡献。     

光照，施肥及刈割对垂穗披碱草生长的影响

以甘南高寒草甸常见牧草垂穗披碱草（Elymus nutans）为研究对象,比较不同光照、肥力条件下,垂穗披碱草对刈割的生长和补偿反应。研究发现,光照和肥力存在交互作用。施肥、中度光照下,未刈割单株垂穗披碱草地上生物量最大,为3．239g。施肥、中度光照下,比较刈割植株地上生物量补偿发现：晚期刈割单株地上生物量的补偿大于早期刈割;刈割强度无显著影响;刈割两次单株地上生物量的补偿最大。施肥、中度光照、晚期、轻度、一次刈割时,单株垂穗披碱草地上生物量累积最大,为4．500g。全光照、未施肥条件下,刈割对单株地上生物量的补偿无影响。就补偿机制而言,植株地上生物量的增加以地下生物量为代价。     

施肥和刈割对青藏高原东部高寒草甸群落生物量补偿效应的影响

通过对青藏高原东部高寒草甸植物群落在4个施肥处理及7个刈割水平下的群落生物量、物种丰富度和补偿指数的比较,分析了该类草甸群落主要特征对施肥及刈割强度差异的响应.结果表明:施肥显著增加了群落生物量,并降低物种丰富度,而刈割处理虽然增加了物种丰富度,但对群落生物量无显著影响;随着施肥量的增加,高寒草甸由莎草科和杂草类物种占优势的群落构成逐渐向禾本科物种占优势的群落构成转变;施肥可提高群落的补偿能力,在不同施肥梯度中,刈割后植物群落补偿能力也不相同;随着施肥量的增加,增加刈割强度,减少刈割次数,发生超补偿的机率增大;可见,施肥是主效应,刈割对群落主要特征的影响随营养水平的不同而发生变化.     

青藏高原东部高寒草甸群落生物量和补偿能力对施肥与刈割的响应

以青藏高原东部高寒草甸群落为研究对象,通过比较了不同施肥条件和不同刈割对群落地上生物量和多样性的影响。结果表明施肥可提高生物量且生物多样性降低,施肥和刈割处理后,施肥效应显著而刈割效应不显著,说明施肥是主效应。实验还发现施肥可提高群落的补偿能力;不同资源梯度的情况下植物群落对刈割处理后补偿作用也不相同,对刈割处理后植物群落补偿能力随资源的升高而增强。当未施肥和施肥30g/m^2时相同强度的1次刈割的补偿能力较相同强度的2次刈割的补偿能力大;当施肥60g/m^2和120g/m^2时相同强度的2次刈割的补偿能力较相同强度的1次刈割的补偿能力大。     

刈割、施肥和浇水对矮嵩草补偿生长的影响

通过对青海海北高寒矮嵩草(Kobresia humilis)草甸进行为期3年的野外控制试验, 研究了刈割(留茬1 cm、3 cm及不刈割)、施肥(2.5 g·m^-2尿素+ 0.6 g·m^-2磷酸二胺、不施肥)和浇水(20.1 kg·m^-2、不浇水)处理对矮嵩草补偿生长(包括分株密度、株高和分株地上生物量)的影响, 及其比叶面积、叶片净光合速率和相对增长率的变化, 探讨矮嵩草补偿生长的机制。研究结果表明: 刈割后, 矮嵩草的补偿生长高度和比叶面积显著降低; 分株密度有增加的趋势, 但会随刈割强度的增加而下降; 株高和生物量的相对增长率随刈割强度的增加而呈上升趋势; 补偿地上生物量在重度刈割处理下最高。施肥能显著增加矮嵩草的补偿高度、分株密度、补偿地上生物量、株高相对增长率、生物量相对增长率、比叶面积和净光合速率; 与不浇水处理相比, 浇水处理对重度刈割处理下的分株地上生物量、密度相对增长率、比叶面积和净光合速率无影响, 而显著降低了中度刈割处理下的补偿高度和株高相对增长率, 提高了不刈割处理下的分株密度和重度刈割处理下的生物量相对增长率。刈割、施肥和浇水处理的交互作用也显示出刈割与施肥对矮嵩草补偿生长具有拮抗效应, 而刈割与浇水具有协同效应。上述结果说明, 矮嵩草在刈割后可通过增加分株密度和相对增长率等途径来提高补偿能力, 弥补在生长高度上出现的低补偿, 而施肥可显著抵消刈割的不利影响, 提高矮嵩草的补偿能力。     

高寒矮嵩草草甸两种主要植物耐牧性的比较

通过野外控制实验,研究了刈割(留茬3 cm、留茬1 cm及不刈割)、施肥(施肥、不施肥)和浇水(浇水、不浇水)处理对高寒草甸矮嵩草(Kobresia humilis)和垂穗披碱草(Elymus nutans)补偿高度、株高相对生长率、比叶面积、叶片净光合速率和地上总生物量的影响.结果表明:2物种的株高和地上总生物量在刈割后均为低补偿响应,但其株高相对生长率显著提高,并均随年份而增加;垂穗披碱草比叶面积、叶片净光合速率和地上总生物量对刈割损伤更加敏感;尽管施肥能显著提高2物种上述各项指标,但在不同处理条件下矮嵩草的耐牧性指数均小于垂穗披碱草;浇水的作用不显著.说明2物种的耐牧性依赖于土壤养分资源获得性,矮嵩草的耐牧性强于垂穗披碱草.     

刈割、施肥和浇水对垂穗披碱草补偿生长的影响

植物的补偿生长特性受放牧强度和生境资源获得性的影响。通过为期2年的野外控制实验,研究了刈割高度(留茬1cm、3cm及不刈割)、施肥(施、不施)和浇水(浇、不浇)处理对垂穗披碱草(Elymus natans)补偿生长的影响,并结合对各处理分株密度、比叶面积、净光合速率和相对生长率的变化研究,探讨了其补偿生长机制。结果表明:刈割后垂穗披碱草分株种群密度显著增加,补偿生长高度显著降低,比叶面积和相对生长率随刈割强度增加而呈上升趋势,叶片净光合速率变化不显著;施肥能显著增加垂穗披碱草的补偿生长高度、比叶面积、叶片净光合速率和相对生长率;浇水处理以及刈割、浇水、施肥处理之间的交互作用均不显著。可见,在刈割条件下,垂穗披碱草具有一定的密度补偿机制,但由于刈割抑制补偿性高生长,导致分株高度出现低补偿。因此,即使刈割后比叶面积和株高相对生长率显著增加,也不一定必然引起株高的超补偿;但施肥可显著提高垂穗披碱草的补偿能力,增加耐牧性,证实了改进后的限制资源模型的预测。     

5种入侵植物补偿反应及其形态可塑性比较

通过模拟刈割试验,研究了5种外来入侵植物(黑麦草、空心莲子草、三叶鬼针草、薇甘菊和胜红蓟)的补偿能力及其形态可塑性。结果表明:多数植物的补偿能力与形态可塑性有一定相关性,刈割和非刈割条件下空心莲子草和薇甘菊都具有较强的形态可塑性,其补偿指数较高,分别为1.65和1.27;多年生黑麦草和三叶鬼针草的形态可塑性在刈割条件下得到增强,补偿指数分别为1.44和1.28;刈割条件下胜红蓟的形态可塑性变化不大或有所降低,其补偿指数最低,为0.82。研究认为,5种植物具有不同的限制其形态可塑性和补偿能力比较理想的刈割时期,薇甘菊攀援生长方式下为30d前和60d后,空心莲子草为60d后,胜红蓟为40～50d,鬼针草为30d前,黑麦草为40～50d。     

苗期刈割伤害对春小麦影响的盆栽实验研究

盆栽实验研究了春小麦在苗期（三叶期）受到不同强度刈割伤害（模拟动物的采食）后的补偿作用。结果表明,在保持田间持水量６０％和９０％两种水分梯度下,春小麦苗期的刈割伤害均可刺激生长,出现生物产量和经济产量的超越补偿,而且受到重度刈割伤害春小麦（Ｅ、Ｆ）的补偿作用大于受到轻度刈割伤害春小麦（Ｂ、Ｄ）的补偿作用。这一超越补偿作用是以耗费较多的水分为基础的。     

Overall grazing tolerance index (overall GTI) is not an ideal predictor for describing a single‐species tolerance to grazing

Plants' pattern of compensatory growth is often used to intuitively estimate their grazing tolerance. However, this tolerance is sometimes measured by the overall grazing tolerance index (overall GTI), which assumes that tolerance is a multivariate linear function of various underlying mechanisms. Because the interaction among mechanisms is not independent, the grazing tolerance expression based on overall GTI may be inconsistent with that based on compensatory growth. Through a manipulative field experiment from 2007 to 2012, we measured the responses of 12 traits of Elymus nutans to clipping under different resource availabilities in an alpine meadow and explored the compensatory aboveground biomass and the overall GTI to assess the possible differences between the two expressions of tolerance. Our results showed that these two expressions of tolerance were completely opposite. The expression based on overall GTI was over‐compensatory and did not vary with clipping and resource availability, while the expression based on compensatory aboveground biomass was under‐compensatory and altered to over‐compensation after fertilization. The over‐expression of highly variable traits with extremely high negative mean GTI to defoliation damage, the influence of random errors contained in traits considered, and the doubling weight of functional redundant traits greatly inflated the overall GTI, which leads to the inconsistency of the two tolerance expressions. This inconsistency is also associated with the different determining mechanisms of the two tolerance expressions. Our data suggest that plants' grazing tolerance is not a multivariate linear function of traits or mechanisms that determine grazing tolerance; the overall GTI is only a measure of traits' variability to defoliation damage. Our findings highlight that the tolerance of E. nutans mainly depends on the response of traits with lower variability to defoliation, and the overall GTI is not an ideal predictor for describing a single‐species tolerance to grazing.     

Interactive effects of clipping and nutrient availability on the compensatory growth of a grass species

Resource availability is an important factor affecting the capacity of compensatory growth after grazing. We performed a greenhouse experiment with Poa bulbosa, a small perennial grass of the Mediterranean and Central Asian grasslands, to test the importance of nutrient availability for compensatory growth after clipping. We also compared the results with predictions of the limited resource model (LRM). Plants were grown at low and high fertilization levels and subjected to a clipping treatment. Contrary to the LMR, we found that in Poa plants compensatory growth occurred under the high fertilization level, while it did not occur under the low level. The LMR predicts a higher tolerance for grazing in the stressful environment. Our plants showed a significant decrease in their relative growth rates (RGR) after clipping. Although the plants allocated a 32–188% greater fraction of the mass to lamina growth after clipping, this greater allocation to the leaves did not fully compensate for the initial reduction in leaf area ratio (LAR). A sensitivity analysis showed for the clipped plants under the high fertilization treatment, that changes in leaf allocation (f _lam) enabled the plants to compensate for a part of the potential loss caused by defoliation. Probably, the increased biomass allocation comes largely from the bulbs. We conclude that the inconsistency of the LRM with our results originates in the lack of compensatory mechanisms in the model. To better understand how environmental conditions affect tolerance to herbivory, the effects of compensatory growth should be taken into account.     

Interactions between herbivory and resource availability on grazing tolerance of Leymus chinensis

Herbivory and resource interact to influence plant regrowth following grazing, but few detailed investigations on grazing tolerance at population levels are available. We conducted two pot experiments along a simulated grazing gradient (0%, 25%, 50% and 75% of shoot removal) at three water or nutrient levels to determine the interaction of resource and herbivory on Leymus chinensis, a perennial, dominant species in the eastern Eurasian steppes. Interactions between water availability and clipping intensity on the relative height growth rate (RHGR) and bud number were significant. Significant interactions between nutrient and clipping on RHGR, total biomass and specific leaf area (SLA) were also found. Total biomass and bud number, showing a unimodal curve along the clipping gradient in resource-rich environments, were highest at light clipping level, suggesting that this species has the plastic compensatory responses from under- to overcompensation. Interactions between herbivory and water or nutrient were opposite to each other. The “cooperative” interactions between water and herbivory magnified the difference in grazing tolerance of L. chinensis between high and low water treatments. The “antagonistic” interactions between nutrient and herbivory, on the other hand, were reflected in the lower tolerance to heavy clipping in the high nutrient than low nutrient treatments. Results partly support the limiting resource model (LRM). A modified and simplified graphic model of the LRM was proposed based on our results. The new LRM clearly demonstrated that “cooperative” interactions between varying water levels and clipping intensities aggravate the detrimental impacts of herbivores on plant growth and reproduction, whereas “antagonistic” interactions between nutrient and grazing alleviate the negative effects of herbivores. Biomass compensation and density compensation were identified as main mechanisms of herbivory tolerance in this clonal species.     

Patterns of growth compensation in eastern white pine (Pinus strobus L.): the influence of herbivory intensity and competitive environments

To investigate complex growth compensation patterns, white pine (Pinus strobus L.) seedlings were clipped to simulate different herbivory levels. Seedlings were growing with different understory competition levels (created through monthly weeding vs no brush control) under a range of overstory canopy closures. Compensation patterns varied for the different growth and size measures. After one growing season, seedlings did not fully compensate for lost biomass regardless of the competitive environments of the seedlings. Although relative height growth was stimulated by light intensity clipping (20-40% of last-year shoots removed), relative diameter growth, total biomass, and biomass growth of seedlings declined sharply with increasing clipping intensity. Likewise, all growth parameters declined with increasing interspecific competition. Results showed that seedlings in highly competitive environments showed smaller growth loss due to clipping than those in competition-free environments, presumably because seedlings experiencing high interspecific competition devoted more energy to maintaining apical dominance and a balanced shoot-root ratio. While competition from canopy trees altered compensatory patterns, competition from understory vegetation only altered the magnitude, but not the patterns, of compensatory growth. We suggest that compensatory growth follows a complex pattern that will vary with the parameters measured, competitive conditions, and clipping intensities. Our results support the assertion that overcompensation may be an adaptation to competitive ability, rather than a response to herbivory itself.     

A test of the compensatory continuum: fertilization increases and below-ground competition decreases the grazing tolerance of tall wormseed mustard (Erysimum strictum)

Contrary to the general expectation, the compensatory continuum hypothesis proposes that grazing may not always affect plant performance adversely. Instead, the effects may vary from negative (undercompensation) to positive (overcompensation), depending on the local availability of resources and the intensity of competition experienced by individual plants. We tested this hypothesis in a common garden experiment by growing tall wormseed mustard, Erysimum strictum, under a factorial design involving simulated grazing (0, 10, or 50% of the main stem clipped), supplemental fertilization and below-ground competition. The results supported the hypothesis. On an average, fertilization increased and competition decreased plant performance. Overcompensation was only observed among the fertilized plants growing free of competition. Simulated grazing increased seed yield 1.6 (10% clipping) and 1.4 times (50% clipping) as compared to unclipped plants when the plants were grown with fertilization and without competition. In contrast, clipping did not significantly increase seed yield in the plants grown without fertilization and/or with competition. The breakage of apical dominance provides a proximate mechanism of these regrowth responses. This is consistent with the fact that most plants (85%) had an unbranched shoot architecture in our study population. However, it is not clear why E. strictum has a relatively unbranched architecture in natural populations. We briefly discuss the alternative ecological factors – competition for light, adaptation to herbivory and optimal timing of flowering as a bet-hedging strategy in monocarpic plants – which might maintain unbranched architecture in this species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Overcompensation and adaptive plasticity of apical dominance in Erysimum strictum (Brassicaceae) in response to simulated browsing and resource availability

In the cases where overcompensation has been observed in monocarpic herbs, overcompensation is associated with an apically dominant shoot architecture of intact plants, increased lateral branching following herbivory, and increased reproductive success as a consequence of damage. The compensatory continuum hypothesis expects overcompensation to be more prevalent in resource rich environments compared to poor environments. This is paradoxical since in resource rich conditions the intact plants should branch most vigorously and hence any further increase in branch number should lead to lower seed yield. An explanation could be that apical dominance is rather insensitive to changes in resource availability, and that overcompensation is possible in conditions where plants experience meristem limitation (due to apical dominance) in relation to available resources. We explored the branching patterns and fitness responses of tall wormseed mustard (Erysimum strictum) to simulated browsing, soil nutrients, and competition in common garden. Competition increased apical dominance and reduced plant fitness whereas fertilization had the reverse effects. Simulated browsing increased lateral branching and had little impact on plant fitness. Fitness overcompensation was observed only among plants grown in competition and in the absence of fertilization – the most resource poor treatment combination in the experiment. The results contradict both with the compensation continuum and the assumption that apical dominance shows no or very little plasticity in relation to growing conditions. Because directional selection gradients on branch number were invariantly positive irrespective of growing conditions, we propose that, in spite of phenotypic plasticity of apical dominance, the plants appear to be meristem rather than resource limited, and that meristem limitation is strongest in conditions where intact plants produce fewest lateral branches. Our results deviate from the compensation continuum because resource availability affected compensation ability more strongly through phenotypic plasticity of shoot architecture rather than via changes in resource availability per se.     

植物耐虫性研究进展

本文简要介绍了植物耐虫性的含义、发生范围、耐虫性的进化过程和遗传特性、耐虫性机理以及影响植物耐虫性表达的非生物和生物因子。植物耐虫性机理的研究涉及光合作用能力变化、同化产物的再分配、内源激素的变化、休眠分生组织的激活和补偿生长、储藏器官的利用、植物物候学和植株株型结构的变化等。研究表明,植物受害后光合作用强度的变化与其耐虫性没有相关性,有些耐虫植物受害后光合作用能力增加,有些植物光合作用强度无明显影响或者下降较少; 害虫取食为害可促进耐虫植物的同化产物得到最大程度利用,能激活耐虫植物的休眠分生组织,产生超补偿作用; 耐虫植物受害部位细胞分裂素含量显著升高; 虫害引起物候学变化小的植物具有较强的耐虫性; 植物的冠层结构、叶形态、根茎比、茎蘖数等植株株型变化与耐虫性有关。影响植物耐虫性表达的因子主要有温度、大气CO₂浓度、土壤营养水平、农用化学物质、植株年龄、害虫分布类型和取食方式、植物共生物等。不同植物在相同温度下对同一种害虫的耐害性差异大,其主要原因可能是由于温度的变化引起同化产物的分配和再分配以及气孔关闭对气体交换和光合作用能力的影响; 生长在高CO₂含量大气中的植物,对害虫的为害有较强耐受性。土壤营养水平对植物耐虫性表达的影响大于温度,增施磷、钾肥可增加植物的耐虫性。聚集分布型害虫为害对植物造成的损失大于随机分布型和均匀分布型害虫,害虫的取食方式、传粉昆虫的活动、植物内生真菌和菌根真菌的感染均影响到植物耐虫性的表达水平。文中最后讨论了植物耐虫性在害虫综合治理中的重要性及应用前景。