Differences in the compensatory growth of two co-occurring grass species in relation to water availability

We compared the potential for compensatory growth of two grass species from the Mongolian steppe that differ in their ability to persist under grazing: the rhizomatous Leymus chinensis and the caespitose Stipa krylovii, and investigated how this ability might be affected by drought. Plants were grown in a greenhouse under wet and dry conditions and subjected to a clipping treatment (biweekly removal of 75–90% of the aerial mass). Leymus exhibited a much stronger compensatory growth after clipping than Stipa. Leymus showed a significant increase in its relative growth rate (RGR) after clipping, while for Stipa RGR was negatively affected. Clipped Leymus plants maintained leaf productivity levels that were similar to undamaged individuals, while leaf-productivity in clipped Stipa dropped to less than half of that of the controls. In Leymus, there was less compensatory growth under dry than under wet conditions, while in Stipa the compensation was increased under drought. This difference probably reflects the fact that Stipa is more drought-tolerant than Leymus. The greater compensatory growth of Leymus compared to Stipa mainly resulted from a greater stimulation of its net assimilation rate (NAR), and its greater capacity to store and reallocate carbohydrates by clipping. The greater increase in NAR was probably the result of a stronger reduction in self-shading, because Leymus shoots were much denser than those of Stipa, which resulted in a higher increase in light penetration to remaining leaves after clipping. The results of this study suggest that the greater ability of Leymus to persist under grazing is the result of its larger capacity for compensatory growth.     

Effects of nutrient supply,light availability and herbivory on the growth of heather and three competing grass species

Grasses are becoming more abundant in areas in NE Scotland which until recently were dominated by heather (Calluna vulgaris). However, it is not clear if grasses are aggressive competitors which are now able to outcompete the dwarf shrub due to changes in environmental factors (such as grazing pressure and increasing nutrient inputs), or just opportunistic invaders, occupying gaps in the canopy which occur when heather reaches the degenerate stage. Experiments in turves and in field plots were carried out in order to investigate the performance of three grass species, Nardus stricta, Deschampsia cespitosa and Deschampsia flexuosa growing in competition with heather. These three species were selected because they differ in their nutrient requirements, palatability to herbivores and tolerance of shading. The grasses were planted in heather canopies of different structure, either turves of heather of different height and age, or moorland plots with or without heavy grazing by sheep and deer. Fertiliser (NPK) was applied to half the experimental plants. The growth of the grass species and the heather in response to the fertiliser and grazing treatments was measured, together with the light levels penetrating the canopy and received by the grass plants.Results indicated that heather was likely to be outcompeted by grasses only when there are gaps in the canopy, resulting either from heavy grazing or from the heather being in the mature or degenerate phase. Fertiliser enhanced plant growth whereas fencing out herbivores led to strong competition for light as the heather canopy closed. It is concluded that grasses require gaps in the canopy to successfully invade heather moorland, or they tend to be shaded out. Thus better management of heather moorlands to maintain a dense canopy structure may help to preserve heather cover even under increasing nutrient inputs.     

Effects of ramet clipping and nutrient availability on growth and biomass allocation of yellow nutsedge

A glasshouse experiment was conducted to examine how the interactions of nutrient availability and partial ramet clipping affect growth, reproduction and biomass allocation of Cyperus esculentus, an invasive sedge. The plants sprouting from tubers were grown at low and high nutrient levels, and were subject either to no clipping, one, two or three clippings, with each clipping cutting half of the existing ramets at soil level. Our results show that nutrient availability and clipping frequency tended to independently affect most of growth, reproduction and biomass allocation parameters of Cyperus esculentus examined in the present study. Increased supply of nutrients led to an increase in plant productivity and its associated traits. All of the traits, except for the number of ramets, displayed a decreasing pattern with increasing clipping frequency, indicating that Cyperus esculentus had undercompensatory responses to ramet clipping. It is likely that the patterns of plants response to clipping are species specific, and depend on morphological characters of species. Its susceptibility to ramet clipping can offer opportunities for controlling this invasive species through mechanical methods such as mowing. Clipping had little effects on biomass allocation; however, root weight fraction increased with increasing clipping frequency. While nutrient availability and clipping frequency had no influence on leaf carbon concentration at harvest, both of them increased leaf nitrogen concentration, and hence reduced leaf C/N ratio.     

Root morphological plasticity and nutrient acquisition of perennial grass species from habitats of different nutrient availability

We studied the root foraging ability and its consequences for the nutrient acquisition of five grass species that differ in relative growth rate and that occur in habitats that differ widely in nutrient availability. Foraging responses were quantified, based on the performance of the plants in homogeneous and heterogeneous soil environments of the same overall nutrient availability. Although all species tended to produce a significantly higher root length density in a nutrient-rich patch, this response was significant only for the faster-growing species. The increased root length density resulted from small, though not significant, changes in root biomass and specific root length. The effectiveness of root proliferation was determined by quantifying the total amount of nutrients (N and P) accumulated by the plants over the course of the experiment. Plants acquired more N in a heterogeneous environment than in a homogeneous environment, although the total nutrient availability was the same. The ability to acquire nutrients (N or P) in the heterogeneous environment was not related to the ability of species to increase root length density in response to local nutrient enrichment. In contrast to other studies, our results suggest that the role of morphological plasticity of roots in acquiring patchily distributed resources is limited. Possible reasons for this discrepancy are discussed. Received: 11 September 1997 / Accepted: 28 February 1998     

Interactive effects of redox intensity and phosphate availability on growth and nutrient relations of Cladium jamaicense (Cyperaceae)

Expansion of Typha domingensis into areas previously dominated by Cladium jamaicense in the Florida Everglades has been linked to anthropogenic phosphorus (P) enrichment and increased hydroperiod. The principal stress factor for plants in flooded soils is biochemical reduction, the intensity of which is measured as redox potential (Eh). The objective of this study was to assess the growth response of C. jamaicense to Eh (-150, +150, and +600 mV) and P availability (10, 80, and 500 μg P/L). Plants were grown hydroponically in a factorial experiment using titanium (Ti(3+)) citrate as an Eh buffer. Treatment effects on growth, biomass partitioning, and tissue nutrients were recorded. Growth approximately doubled in response to a 50-fold increase in P availability. Low redox significantly reduced growth and tissue P concentration. While plant P concentrations increased 20-fold between the 10 and 500 μg P/L treatments, P concentrations were 50-100% higher at +600 mV than at -150 mV within each phosphate level. At high Eh, C. jamaicense appears well adapted to low nutrient environments because of its low P requirement and high retention of acquired P. However, at low Eh the ability to acquire or conserve acquired P decreases and as a consequence, higher phosphate levels are required to sustain growth. Findings of this study indicate that young C. jamaicense exhibits low tolerance to strongly reducing conditions when phosphate is scarce.     

放牧格局和生境资源对矮嵩草（Kobresia humilis）分株生物量分配和补偿性生长的影响

为研究密集型克隆植物对放牧扰动和生境资源变化的生物量分配和补偿生长响应特性,验证克隆植物的觅食模型和3个有关植物个体补偿反应的假说(①放牧优化假说,grazing optimization hypothesis,GOH;②反应连续谱假说,continuum of responses hypothesis,CRH;③增长率模型,growth rate model,GRM),在具有不同放牧利用格局和土壤养分水平的高寒矮嵩草草甸(Ⅰ.畜圈草地:重度放牧、资源丰富;Ⅱ.牧道草地:中度放牧、资源贫乏;Ⅲ.封育草地:不放牧、资源贫乏)中通过设置扣笼/无扣笼样方对其建群种矮嵩草(Kobresia humilis)进行了研究.结果表明不论当年解除家畜放牧与否,春季采摘率越高(畜圈草地),分株生物量向生长的投入越少,向贮藏器官的投入越多.繁殖分配在中度采摘下最高(牧道草地),扣笼内外分株各部分的生物量分配无差异.矮嵩草分株在中度采摘×资源贫乏条件下产生了超补偿响应,在重度采摘×资源丰富条件下为等量补偿,重度采摘导致分株密度显著减少.生物量分配格局与觅食模型的预测不符.补偿生长响应特性证实了GOH和GRM的预测,但与CRH的预测不符.这说明在研究地区放牧扰动格局对克隆植物矮嵩草分株的生物量分配和补偿生长具有重要影响,适度放牧利用更利于引起超补偿,而重度利用可能会对该种群的长期保持产生不利影响.     

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。     

Coping with low nutrient availability and inundation: root growth responses of three halophytic grass species from different elevations along a flooding gradient

We describe the responses of three halophytic grass species that dominate the low (Spartina anglica), middle (Puccinellia maritima) and high (Elymus pycnanthus) parts of a salt marsh, to soil conditions that are believed to favour contrasting root-growth strategies. Our hypotheses were: (1) individual lateral root length is enhanced by N limitations in the soil but restricted by oxygen limitations, (2) the density of root branching within a species is inversely related to the length of the lateral roots, and (3) species from high elevations (i.e. the driest parts of a marsh) are the most responsive to changing soil conditions. Plant growth responses and soil parameters showed that the contrasting but uniformly applied soil treatments were effective. All three species showed a small but significant shift towards a finer root diameter distribution when N was limiting, partly because of the finer diameters of the laterals (Elymus and Spartina) and partly because of increased length of individual 1st-order laterals (Elymus and Puccinellia). The increased length of the 1st-order laterals of Elymus and Puccinellia grown under low N indicates that the first part of hypothesis 1 may be true. However, lack of effect of flooding and reduced soil conditions lead us to reject the second part of hypothesis 1. Hypothesis 2 was rejected for these three halophytes, as the branch density of 1st- and 2nd-order laterals appears to be controlled by other factors than length of individual laterals. Hypothesis 3 may be true for specific root characteristics (e.g. length of individual 1st-order laterals), but cannot be generalised (e.g. branch density and topological index). In conclusion, the present data on root growth in contrasting but homogeneous soil conditions indicate that morphological responsiveness of the root systems of these halophytic grass species is limited, regardless of their location along the elevational gradient.     

Interactive effects of plant-available soil silicon and herbivory on competition between two grass species

Background and Aims

The herbivore defence system of true grasses (Poaceae) is predominantly based on silicon that is taken up from the soil and deposited in the leaves in the form of abrasive phytoliths. Silicon uptake mechanisms can be both passive and active, with the latter suggesting that there is an energetic cost to silicon uptake. This study assessed the effects of plant-available soil silicon and herbivory on the competitive interactions between the grasses Poa annua, a species that has previously been reported to accumulate only small amounts of silicon, and Lolium perenne, a high silicon accumulator.

Methods

Plants were grown in mono- and mixed cultures under greenhouse conditions. Plant-available soil silicon levels were manipulated by adding silicon to the soil in the form of sodium silicate. Subsets of mixed culture pots were exposed to above-ground herbivory by desert locusts (Schistocerca gregaria).

Key Results

In the absence of herbivory, silicon addition increased biomass of P. annua but decreased biomass of L. perenne. Silicon addition increased foliar silicon concentrations of both grass species >4-fold. Under low soil-silicon availability the herbivores removed more leaf biomass from L. perenne than from P. annua, whereas under high silicon availability the reverse was true. Consequently, herbivory shifted the competitive balance between the two grass species, with the outcome depending on the availability of soil silicon.

Conclusions

It is concluded that a complex interplay between herbivore abundance, growth–defence trade-offs and the availability of soil silicon in the grasses'' local environment affects the outcome of inter-specific competition, and so has the potential to impact on plant community structure.     

The effects of water,nutrient availability and their interaction on the growth,morphology and physiology of two poplar species

We exposed cuttings of two poplar species, Populus cathayana Rehder and Populus przewalskii Maximowicz, from Sect. Tacamahaca Spach to two watering regimes (well-watered and water-stressed conditions) and to two nutrient regimes (with or without fertilization) in a greenhouse to determine how fertilization affects the growth, morphology and physiology of poplars under different water conditions. Under stress conditions, changes in early growth and dry matter allocation, and decrease in gas exchange and the related functions are usually observed. Moreover, the measurement of carbon isotope composition (δ¹³C) provides an integrated measurement of water use efficiency. And abscisic acid (ABA) is a phytohormone which plays a prominent role in various physiological and biochemical processes related to environmental stresses. So we determine these characteristics and related parameters, and our results showed the following: (1) Fertilization promoted the growth of poplars under well-watered conditions, while under water-stressed conditions its effect on growth was negative. (2) Fertilization increased δ¹³C, total N concentration, chlorophyll a/b and intrinsic efficiency of photosystem II (Fv/Fm) but decreased relative water content of leaves, stomatal conductance, transpiration rate and C/N ratio under both well-watered and water-stressed conditions. (3) Fertilization appeared to increase net photosynthesis rate and decrease ABA content under well-watered conditions, while it decreased net photosynthesis rate and increased ABA content under water-stressed conditions. Moreover, compared to P. cathayana, collected from a lower altitude region, P. przewalskii, collected from a high-altitude region, has a slower growth rate and stronger adaptability to drought stress, which perhaps resulted from its chronic adaptability to the low water availability of high-altitude region; but to the nutrient stress, there was no difference between the two species.     

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。     

Interactive effects of elevated summer temperature,nutrient availability,and irradiance on growth and chemical compositions of juvenile kelp,Eisenia bicyclis

Previous experiments with a factorial design have revealed the effects of several environmental factors on species performance and their interactions, which indicate synergistic or antagonistic effects. Temperature, nutrient availability, and irradiance are well‐known environmental factors that affect the growth and chemical composition of brown algae. However, relatively few studies have tested their combined effect on brown algal growth and chemical composition using a three‐way factorial design. We conducted a culture experiment to test the combined effects of elevated summer temperatures (23 and 26°C), irradiance (180 and 30 μmol photon m^?2 s^?1), and nutrient availability (enriched and non‐enriched seawater) on four relative growth rates (RGRs; based on wet weight, blade width, length, and area) and three chemical compositions (including carbon, nitrogen, and phlorotannin content) in juvenile sporophytes of the kelp Eisenia bicyclis. RGR based on blade width was the most sensitive to abiotic factors among all RGRs. A significant interaction between temperature and nutrient availability on this RGR suggested that the negative effect of elevated temperature was antagonized by a reduction in nutrient availability. Similarly, the positive effect of elevated irradiance on carbon content was synergized by reduced nutrient availability. Moreover, the negative effect of increased irradiance on nitrogen content was antagonized by elevated temperature in nutrient‐enriched treatments, but not in non‐enriched treatments. The content of carbon‐based phlorotannins increased with reduced nutrient availability but not with elevated irradiance. These results suggest that these abiotic factors have complex interactions on the growth and chemical composition of this species.     

The effect of clipping frequency on the competitive interaction between two perennial grass species

Summary The effect of clipping frequency on competition between Lolium perenne and Agrostis tenuis was investigated. The yield of clippings of both species increased and then declined during the 12-week period of the experiment, but the clip yield of Lolium was always significantly greater than that of Agrostis. Lolium was clearly the better competitor in unclipped controls. The proportion of the biomass contributed to the mixture by Agrostis increased as the interval between clips decreased. Tiller production was unaffected by increased clipping frequency in Lolium but was increased in Agrostis. Total yield was much more drastically reduced by frequent clipping in Lolium than in Agrostis, where yield was practically unaffected by wide variations in clipping frequency.These results are in agreement with the field distributions of the two species. They also suggest that differences in height and response to clipping are likely to confound any attempt to monitor the progress of competition experiments by measuring the yield of clippings.     

Interactive effects of protein and energy intake on nutrient partitioning and growth in Nile tilapia

Studies of fish growth response to changes in dietary protein and energy content are often conducted with fish fed to apparent satiation or at fixed percentages of their body mass. Such designs result in simultaneous changes in protein and non-protein energy intake, thereby failing to distinguish their separate effects on nutrient partitioning and growth. The present study was designed to address this limitation and test the existence of distinct protein- and non-protein energy-dependent growth phases in Nile tilapia (Oreochromis niloticus). All-male Nile tilapia (63 g, SD = 1.3) were subjected to an 8 × 2 factorial design consisting of eight levels of digestible protein (DP) intake (0.44–1.25 g/day) and two levels of non-protein digestible energy (NPDE) intake (16.0 and 22.4 kJ/day). Fish (n = 960) were housed in 60-litre tanks with two replicates per treatment and hand-fed twice a day for 42 days. Nutrient balances were calculated from changes in body mass, analysed body composition and digestible nutrient intake. Linear regression models were compared to linear-plateau regression models to determine whether protein gain followed distinct protein- and non-protein energy-dependent phases or not. Body mass gain increased linearly with increasing DP intake and was significantly higher (2.6 vs 2.3 g/d, P < 0.05) in fish receiving a high NPDE intake. This increase mainly reflected a higher mean fat gain (0.29 vs 0.20 g/d) rather than a higher protein gain (0.42 vs 0.39 g/d) in fish fed a high vs low level of NPDE intake. The comparison of linear and linear-plateau models did not give clear support for the presence of distinct protein and non-protein energy-dependent phases in protein gain. These results indicate that non-protein energy intake has a modest protein-sparing potential, and that protein gain is simultaneously limited by protein and energy intake in Nile tilapia.     

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。     

Fire and nutrient gradient effects on the sapling ecology of four Acacia species in the presence of grass competition

The variation among tree species in their sapling responses to disturbances such as fire, herbivory and grass competition results in different probabilities of recruitment into savanna communities. A sapling??s ability to persist under these disturbances depends on its investment in growth, defences and stored reserves, based on soil available nutrients. We examined the survival and investments in growth and defence of the saplings of four Acacia species, in the presence of competing grasses, under varying soil nutrient levels. We also have evaluated the effects of fire on these saplings. We performed the study in a greenhouse at the UKZN Botanical Garden, Pietermaritzburg, South Africa. Nutrients had no significant effect on the survival of tree saplings regardless of fire treatment. Only 22.5?% saplings survived after fire treatment compared with 84?% of control saplings. Acacia sieberiana showed the highest resprout survival, while A. karroo did not survive the fire. Growth in A. nigrescens increased only at intermediate levels (0.5N, 1N) of the fertiliser provided. Condensed tannins in A. karroo also decreased at an intermediate level (1N) of fertiliser. Resprouts grew faster in stem height than in diameter and did not invest in chemical defences. No treatment had an effect on the proportion of root and shoot dry biomass. In conclusion, African acacias are not dependent on soil nutrients for their survival and differ in their ability to resprout after fire. Intermediate levels of nutrient availability are beneficial to sapling development as opposed to zero and high nutrient levels, due to nutrient-grass dynamics. The benefits of post-fire grass removal and an increase in nutrients can be effectively utilised by only a few species of acacias.     

Effect of species on dune grass growth

Observations of experimental dunes made over a period of nine years indicate differences in utilizing three different dune species along the North Carolina coast and in the type of dunes produced by them.Ammophila is superior in ease of establishment and rate of sand accumulation but is shortlived. It produces a gently sloping dune.Uniola is difficult to propagate but is an excellent dune builder and eventually dominates the foredune. Grown alone, it produces a steep dune front.Panicum is an excellent companion plant to bothAmmophila andUniola. It is easy to propagate, relatively free of pests and produces dunes intermediate in shape, betweenAmmophila andUniola. Plantings of mixtures of two or more of these species should greatly improve long-term dune stability compared with either planted alone.Presented at the Seventh International Biometeorological Congress, 17–23 August 1975, College Park, Maryland, USA     

Interactive effects of food availability and aerial body temperature on the survival of two intertidal Mytilus species

Intertidal organisms must episodically contend with the rigors of both the terrestrial and the marine environments. While body temperatures during high tide are driven primarily by water temperature, aerial body temperatures are driven by multiple environmental factors such that temperature of an organism during low tide is usually quite different from air temperature. Thus, whereas decades of research have investigated the effects of water temperature on intertidal species, considerably less is known about the physiological impacts of temperature during aerial exposure at low tide, especially with regard to the interaction of aerial body temperature with other stressors. We examined the interactive effects of aerial body temperature and food supply on the survival of two intertidal blue mussels, Mytilus galloprovincialis and Mytilus trossulus. Survival was monitored for nine weeks using a simulated tidal cycle, with two levels of food and three levels of aerial body temperature (30, 25, and 20 °C). Decreased food supply significantly reduced the survival of mussels, but only under the 30 °C treatment. In the other two thermal regimes there were no significant effect of food on survival. When aerial body temperatures are high, food availability may have a greater effect on intertidal organisms. Decreases in ocean productivity have been linked to increased in ocean temperatures, thus intertidal organisms may become more susceptible to thermal stress as climates shift.