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     

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 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.     

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.     

Interactive effects of nutrient and mechanical stresses on plant morphology

BACKGROUND AND AIMS: Plant species frequently encounter multiple stresses under natural conditions, and the way they cope with these stresses is a major determinant of their ecological breadth. The way mechanical (e.g. wind, current) and resource stresses act simultaneously on plant morphological traits has been poorly addressed, even if both stresses often interact. This paper aims to assess whether hydraulic stress affects plant morphology in the same way at different nutrient levels. METHODS: An examination was made of morphological variations of an aquatic plant species growing under four hydraulic stress (flow velocity) gradients located in four habitats distributed along a nutrient gradient. Morphological traits covering plant size, dry mass allocation, organ water content and foliage architecture were measured. KEY RESULTS: Significant interactive effects of flow velocity and nutrient level were observed for all morphological traits. In particular, increased flow velocity resulted in size reductions under low nutrient conditions, suggesting an adaptive response to flow stress (escape strategy). On the other hand, moderate increases in flow velocity resulted in increased size under high nutrient conditions, possibly related to an inevitable growth response to a higher nutrient supply induced by water renewal at the plant surface. For some traits (e.g. dry mass allocation), a consistent sense of variation as a result of increasing flow velocity was observed, but the amount of variation was either reduced or amplified under nutrient-rich compared with nutrient-poor conditions, depending on the traits considered. CONCLUSIONS: These results suggest that, for a given species, a stress factor may result, in contrasting patterns and hence strategies, depending on a second stress factor. Such results emphasize the relevance of studies on plant responses to multiple stresses for understanding the actual ecological breadth of species.     

Additive effects of genotype,nutrient availability and type of tissue damage on the compensatory response of Salix planifolia ssp. planifolia to simulated herbivory

Plant responses to herbivory include tolerance (i.e. compensatory growth) and defense. Several factors influence the tolerance of a plant following herbivory, including plant genetic identity, site nutrient availability, and previous and/or concurrent herbivory. We studied the effects of these factors on the compensatory response of Salix planifolia ssp. planifolia, a shrub species common in the boreal and subarctic regions of North America. We cloned several genets of S. planifolia and submitted them to simulated root and/or leaf herbivory while varying the nutrient availability. Simulated leaf herbivory was more detrimental to the plant than simulated root herbivory, reducing both above- and below-ground tissue production. Leaf demography was unaffected by either simulated herbivory treatment. There was some compensatory growth following simulated leaf and root herbivory, but only the root compartment responded to increased nutrient availability. Simulated leaf herbivory increased leaf transpiration and reduced stomatal resistance, suggesting increased carbon fixation. The unexpected finding of the experiment was the absence of interactions among factors (genotype, nutrient availability and type of tissue damage) on the compensatory response of S. planifolia. These factors thus have additive effects on the species' compensatory ability.     

Interactive effects of soil nutrient heterogeneity and belowground herbivory on the growth of plants with different root foraging traits

Aims

Plants with precise root foraging patterns can proliferate roots preferentially in nutrient-rich soil patches. When nutrients are distributed heterogeneously, this trait is often competitively advantageous in pot experiments but not field experiments. We hypothesized that this difference is due to belowground herbivory under field conditions.

Methods

We performed pot experiments using seedlings of Lolium perenne (a more precise root foraging species) and Plantago lanceolata (a less precise root foraging species). The experiment had a two-way factorial randomized block design, with nutrient distribution pattern (homogeneous or heterogeneous) and belowground herbivore (present or absent) as the two factors. Each pot contained one seedling of each species.

Results

With no herbivore present, plant biomass was smaller in the heterogeneous nutrient treatment than in the homogeneous treatment in P. lanceolata, but not in L. perenne. Under homogeneous nutrient distribution, plant biomass was lower in both species with a herbivore present than with no herbivore. Under heterogeneous nutrient distribution, biomass reduction due to herbivory occurred only in L. perenne.

Conclusions

Roots of the precise root foraging species were grazed more under the heterogeneous nutrient distribution, suggesting that the herbivore more efficiently foraged for roots in nutrient-rich soil patches.     

Host genotype overrides endophyte infection effects on growth,physiology, and nutrient content of a native grass,Achnatherum sibiricum

The effect of infection by the fungal endophyte Neotyphodium, host genotype, and their interaction on growth and physiology, as well as photosynthesis, was investigated in the native grass Achnatherum sibiricum. We artificially inoculated the endophyte into mature tillers of endophyte-free A. sibiricum. Plants were clipped to 5 cm height after recording growth traits, and analyzed for total nonstructural carbohydrates (TNC %), the percentage of nitrogen (N %), and carbon (C %) in leaves before and after clipping. In our study, the prominent host genotype–endophyte infection interactions detected in A. sibiricum indicates that, for many growth and storage traits, endophyte infection can impact a little change. However, there is no overriding consistently positive effect of the endophyte on growth or storage in A. sibiricum before or after clipping. Our study showed that the interaction between endophyte and host grasses was highly contingent on plant genotypes. We found host genotype overrode fungal endophyte infection in influencing tiller number and photosynthetic properties of A. sibiricum before clipping. After clipping, host genotype accounted for more of the variation in regrowth and above-ground biomass of A. sibiricum than endophyte infection. Our study showed that host genotype affected the total nonstructural carbohydrates of A. sibiricum before and after clipping, whereas endophyte infection increased the carbon content after clipping. Genotype by infection interactions for plant height, leaf mass, total nonstructural carbohydrates, and photosynthetic characteristics indicated genotype-specific effects of endophytes on A. sibiricum physiology and photosynthetic capacity. The host genotype–endophyte infection interactions detected in A. sibiricum suggest that host genotype overrides fungal endophyte infection on growth, physiology, and nutrient content of this native grass. In contrast, endophyte effects did not appear to positively affect growth, physiology, or photosynthetic capacity before or after clipping.     

Interacting determinants of specific leaf area in 22 herbaceous species: effects of irradiance and nutrient availability

We measured specific leaf area (SLA) and six of its determinants (the thickness of lamina, mesophyll, epidermis, mid-vein and mid-vein support tissues and leaf water content) in a collection of 22 herbaceous species grown in factorial combinations of high μ 1100 (mol m^–2 s^–1) and low (200) irradiance crossed with high (1 : 1) and low (1 : 6 dilution) concentrations of a modified Hoagland hydroponic solution. SLA increased with both decreasing irradiance and with increasing nutrient availability but there was a strong interaction between the two. Lamina and mesophyll thickness both increased with increasing irradiance and nutrient availability without any interaction. The experimental treatments had complicated effects on mid-vein thickness and its support tissues. Leaf water content (a measure of leaf tissue density) decreased with increasing irradiance levels and with decreasing nutrient supply, but with an interaction between the two treatments. Changes in nutrient supply had no effect on SLA at high irradiance because leaf thickness and leaf tissue density changed in a compensatory way. A path analysis revealed that each of the components affected SLA when the others were statistically controlled but the strengths of the effects of mesophyll thickness, mid-vein thickness and water content differed between treatment groups. The effect of epidermal thickness on SLA was constant across environments and it showed no significant covariation with the other determinants. There was significant covariation between mesophyll thickness, mid-vein thickness and water content and this covariation was constant across the treatment groups.     

Interacting effects of elevated CO2, nutrient availability and plant species on a generalist invertebrate herbivore

By affecting plant growth and phytochemistry elevated CO₂ may have indirect effects on the performance of herbivores. These effects show considerable variability across studies and may depend on nutrient availability, the carbon/nutrient‐balance in plant tissues and the secondary metabolism of plants. We studied the responses to elevated CO₂ and different nutrient availability of 12 herbaceous plant species differing in their investment into secondary compounds. Caterpillars of the generalist herbivore Spodoptera littoralis were reared on the leaves produced and their consumption and growth rates analysed. Elevated CO₂ resulted in a similar increase of biomass in all plant species, whereas the positive effect of fertilization varied among plant species. Specific leaf weight was influenced by elevated CO₂, but the effect depended on nutrient level and identity of plant species. Elevated CO₂ increased the C/N ratio of the leaves of most species. Caterpillars consumed more leaf material when plants were grown under elevated CO₂ and low nutrients. This indicates compensatory feeding due to lower tissue quality. However, the effects of elevated CO₂, nutrient availability and plant species identity on leaf consumption interacted. Both the effects of CO₂ and nutrient availability on the relative growth rate of the herbivore depended on the plant species. The feeding rate of S. littoralis on plant species that do not produce nitrogen‐containing secondary compounds (NCSC) was higher under low nutrient availability. In contrast, in plants producing NCSC nutrient availability had no effect on the feeding rate. This suggests that compensatory feeding in response to low nutrient contents may not be possible if plants produce NCSC. We conclude that elevated CO₂ causes species‐specific changes in the quality of plant tissues and consequently in changes in the preferences of herbivores for plant species. This could result in changes in plant community composition.