Production and nitrogen responses of the African dwarf shrub Indigofera spinosa to defoliation and water limitation

Summary The dwarf shrub Indigofera spinosa Forsk. (Papilionacea), a native forage species of arid Northwest Kenya, was propogated from seed, grown in a controlled environment, and subjected to three treatments of defoliation and watering frequencies in a factorial experimental design. Biomass production and nitrogen accumulation in tissue components were measured to determine defoliation responses in a water-limited environment. We hypothesized that plants would maintain biomass and nitrogen flows despite removal of aboveground meristems and tissues by defoliation. Principal experimental results included a slight reduction (11%; P=0.08) of total biomass production by clipping ca. 1/3 or 2/3 of new leaves and stems and all apical meristems every month. Total aboveground production was not affected by clipping, while final root biomass was reduced 17% by the 2/3 clipping. The least water stressed plants were affected most negatively by defoliation, and the unclipped plants responded more negatively to greater water limitation. Plants achieved partial biomass compensation through alterations in shoot activity and continued allocation of photosynthate to roots. A smaller fraction of leaf production was directed to litter in clipped plants although clipping only removed the youngest tissues, suggesting that clipping increased leaf longevity. In turn, each leaf probably contributed a greater total quantity of photosynthate. Photosynthetic rates were also likely to have been increased by clipping water-stressed plants. In contrast to biomass, plants overcompensated for nitrogen lost to defoliation. Total nitrogen uptake by individual plants was stimulated by defoliation, as there was more total nitrogen in leaves and stems. Increased nitrogen uptake was achieved by clipping stimulation of total uptake per unit of root rather than of total root mass.     

Intraspecific variation in the response of Themeda triandra to defoliation: the effect of time of recovery and growth rates on compensatory growth

Summary The response to a single defoliation was studied on three clones of Themeda triandra collected in the short, mid, and tall grassland regions of the Serengeti National Park (Tanzania). These sites represent a gradient of decreasing grazing intensity. Growth, allocation pattern, and several morphometric traits were monitored during an 80-day period. Clipped plants of the short and medium clones fully compensated for the reduction of biomass, while plants of the tall clone showed overcompensation. During the first two weeks after clipping, clipped plants showed lower relative growth rates than unclipped ones, whereas the opposite was observed later on. Clipped plants compensated for the removal of leaf area by producing new leaves with lower specific weights and higher nitrogen content. They also produced more, smaller tillers. Although clipped plants mobilized nonstructural carbohydrates from roots and crowns, this did not account for a significant amount of growth. Relative growth rates of unclipped plants of the short clone were higher. The relative growth rate of the short clone diminished less after clipping, but also exhibited the lowest increase later. The tall clone was the most negatively affected early, but showed the highest compensation later. Compared to the other clones, the short ecotype showed many of the characteristics that defoliation induced in each individual of any clone: higher allocation to leaf area production, higher relative growth rate, higher number but smaller size of tillers, and lower leaf specific weights.     

The effects of clipping and fertilization on nitrogen nutrition and allocation by mycorrhizal and nonmycorrhizal Panicum coloratum L., a C4 grass

Summary Mycorrhizal and nonmycorrhizal plants of Panicum coloratum L. were grown in a factorial treatment design under two nitrogen levels and two clipping heights with an unclipped control. The nitrogen concentration in different plant components was determined following 9 weeks of growth under experimental conditions. Mycorrhizal infection increased green leaf and sheath nitrogen concentration by a relatively small, but significant percentage and had no effect on nitrogen allocation to the various plant components. Clipping increased leaf nitrogen concentration but inhibited growth to the extent that, when compared with the unclipped controls, less nitrogen remained in residual plant biomass with up to half of the total nitrogen allocated to offtake (the material removed by clipping). Plants receiving the higher nitrogen fertilization had higher tissue concentration of N and more N allocated to above-ground living tissues. Mycorrhizal infection interacted with clipping height and also with N availability significantly. Infection was unable to ameliorate the negative effects of the most severe clipping regime and of the low nitrogen availability on leaf and sheath N content. This is possibly due to mycorrhizal demand for carbohydrates competing with the carbohydrate requirement of roots for nitrogen uptake.     

The interaction of defoliation and nutrient uptake in Sporobolus kentrophyllus,a short-grass species from the serengeti plains

Summary Sporobolus kentrophyllus, a grazing-tolerant C₄ grass from the southeastern Serengeti Plains, was grown in solution culture to examine the effects of clipping on the uptake, preference and subsequent transport of varying nitrogen forms. Clipping reduced offtake mass, crown mass ane root mass, resulting in a 58% decline in plant mass. Proportional biomass allocation to roots decreased with clipping, while tillering rates increased. Clipping also increased the nitrogen concentrations of all tissues, and plant nitrogen uptake (nitrogen accumulated throughout the experiment per gram root). The ¹⁵N concentrations (% atom excess) of all tissues were higher in clipped compared with unclipped plants, and the average ¹⁵N uptake rate of clipped plants was twice that of unclipped plants. The relative ¹⁵N allocation to aboveground mass, a measure of canopy sink strength, was higher in clipped plants. Plants fed ¹⁵N-ammonium or ¹⁵N-nitrate during the ¹⁵N pulse experiment had greater ¹⁵N tissue concentrations compared with urea-fed plants, and ¹⁵N uptake rates were higher in ammonium-fed and nitrate-fed plants, compared with urea-fed plants. The relative magnitudes of these differences were higher when plants were clipped. Clipped plants had higher uptake rates for potassium, phosphorus and sodium, while differences between clipping treatments for calcium, iron, and magnesium were indistinguishable. Rapid uptake rates for species on the southeastern Serengeti plains, particularly during grazing periods, have important implications for nutrient cycling in this system.     

Biomass and mineral element responses of a Serengeti short-grass species to nitrogen supply and defoliation: compensation requires a critical [N]

Large mammalian herbivores in grassland ecosystems influence plant growth dynamics in many ways, including the removal of plant biomass and the return of nutrients to the soil. A 10-week growth chamber experiment examined the responses of Sporobolus kentrophyllus from the heavily grazed short-grass plains of Serengeti National Park, Tanzania, to simulated grazing and varying nitrogen nutrition. Plants were subjected to two clipping treatments (clipped and unclipped) and five nitrogen levels (weekly applications at levels equivalent to 0, 1, 5, 10, and 40 g N m⁻²), the highest being equivalent to a urine hit. Tiller and stolon production were measured weekly. Total biomass at harvest was partitioned by plant organ and analyzed for nitrogen and mineral element composition. Tiller and stolon production reached a peak at 3–5 weeks in unclipped plants, then declined drastically, but tiller number increased continually in clipped plants; this differential effect was enhanced at higher N levels. Total plant production increased substantially with N supply, was dominated by aboveground production, and was similar in clipped and unclipped plants, except at high nitrogen levels where clipped plants produced more. Much of the standing biomass of unclipped plants was standing dead and stem; most of the standing biomass of clipped plants was live leaf with clipped plants having significantly more leaf than unclipped plants. However, leaf nitrogen was stimulated by clipping only in plants receiving levels of N application above 1 g N m⁻² which corresponded to a tissue concentration of 2.5% N. Leaf N concentration was lower in unclipped plants and increased with level of N. Aboveground N and mineral concentrations were consistently greater than belowground levels and while clipping commonly promoted aboveground concentrations, it generally diminished those belowground. In general, clipped plants exhibited increased leaf elemental concentrations of K, P, and Mg. Concentrations of B, Ca, K, Mg, and Zn increased with the level of N. No evidence was found that the much greater growth associated with higher N levels diminished the concentration of any other nutrient and that clipping coupled with N fertilization increased the total mineral content available in leaf tissue. The results suggest that plants can (1) compensate for leaf removal, but only when N is above a critical point (tissue [N] 2.8%) and (2) grazing coupled with N fertilization can increase the quality and quantity of tissue available for herbivore removal. Received: 25 August 1997 / Accepted: 14 April 1998     

Compensatory growth responses to clipping defoliation in Leymus chinensis (Poaceae) under nutrient addition and water deficiency conditions

Compensatory growth responses of Leymus chinensis, a dominant species in Inner Mongolia steppe, to clipping defoliation were evaluated in a pot-cultivated experiment under different nutrient (N and P) and water availability conditions. Leymus chinensis exhibited over-compensatory growth at the light and moderate clipping intensities (20% and 40% aerial mass removed) with a greater accumulated aboveground biomass, higher relative growth rate (RGR), more rhizomatic tillers and a stimulation of compensatory photosynthesis to the remnant leaves as compared with those of the unclipped plants. Intense clipping (80% aerial mass removed), which removed most of the aboveground tissues, greatly reduced the growth of aboveground biomass in comparison with that of the unclipped plants. Nitrogen addition only slightly improved the biomass production and RGR in light and moderately clipped plants, and it did not allow plants in the intense clipping condition to over-compensate. Phosphorus addition had no obvious influences on the growth and physiological responses to clipping defoliation. These results indicated that nutrient addition could not compensate for the negative effects of severe clipping on the defoliated grass. On the other hand, there were no distinct positive responses under water deficiency condition for L. chinensis at all clipping intensities with a significant reduction of aboveground and belowground biomass, lower RGR, fewer rhizomatic tillers, and a lower net photosynthetic rate than other wet treatments. Additionally, the chlorophyll contents of remnant leaves gradually increased with the increase of clipping intensities in each treatment. In conclusion, although L. chinensis could compensate for tissues removal by some morphological and physiological responses, intense clipping and drought can result in a significant decrease of biomass and growth rate, even under enriched nutrition conditions.     

Changes of Total Nonstructural Carbohydrates of Kobresia humilis in Alpine Meadow

The seasonal variation of total nonstructural carbohydrates (TNC) concentration in crowns and roots of Kobresia humilis were determined during 1993～ 1994. The effects of defoliation on TNC concentration and the relationship between the aboveground growth and TNC concentration in crown were analysed. TNC were mostly stored in crowns exhibiting great seasonal fluctuations with narrow V-shaped cycle. TNC concentration of crown varied inversely with the net increased of leaf number in the entire growing season, but it dependedlargely on the dead to survival leaf ratio in the early growth season, with a constant of 20.59% at a ratio of 0.16, either decreased or increased when the ratio was greater or less than 0.16 respectively. In the later, there was a significant positive correlation between the TNC concentration of crowns and the cumulative leaves. However, in mid-season, no significant correlation was found between the aboveground growth and the TNC concentration of crowns. The reserve pools (concentration X crown biomass) had no significant differences among various grazing treatments. Intense clipping at two phenological stages caused reduction of reserves during regeneration, but at the end of the growth season, there was no significant difference between clipped and unclipped plants, as well as between clipping intensities.TNC concentration of crowns and roots significantly decreased following early heavy clipping, but it recovered rapidly in crowns. Clipping at mid-stage affected greatly on TNC con-centration of roots. Clipping influenced more on TNC concentration of roots but grazing had more influence on that of crowns.     

Defoliation-induced enhancement of total aboveground nitrogen yield of grasses

We conducted an experiment in a northern mixed-grass prairie at Wind Cave National Park, South Dakota, USA to evaluate the effect of defoliation frequency on aboveground net primary production (ANPP), shoot nitrogen concentration, and aboveground N yield of graminoids. ANPP was significantly reduced at weekly and biweekly defoliation frequencies, but unaffected relative to unclipped controls at monthly and bimonthly frequencies. By contrast, clipping at all frequencies increased shoot N concentration above that of controls, and this increase was greatest at monthly or more frequent defoliations. Total aboveground N yield and potential N yield to grazers were greatest at intermediate (bimonthly to biweekly) frequencies. We suggest that grazers may maximize their nutritional status in this system by periodically regrazing areas at frequencies near the approximately monthly optimum that we observed.     

Nitrogen allocation in Mojave Desert winter annuals

Summary Nitrogen contents and allocation were examined in winter annuals at two Mojave Desert sites near Boulder City, Nevada. Application of 10 g N m^-2 as NH₄NO₃ increased production 0- to 7-fold in species growing on a sandy soil (an Entisol) but fertilizer had no effect on plants on an alluvium (an Aridisol). Tissue nitrogen comprised 0.09–3.5% of dry weight with the lowest concentrations found in vegetative organs of nitrogen-responsive plants. During development, nitrogenpoor species showed only minor changes in nitrogen concentration and allocation compared with more nitrogen-rich species. Maximum reproductive nitrogen allocation varied among species from 43 to 67%, while reproductive biomass allocation was 31 to 51%. Fertilizer increased reproductive biomass allocation by 7 to 16%, reproductive nitrogen concentrations by 120 to 260%, and eaf and root nitrogen concentrations by 200 to 615% in nitrogen-deficient plants. Nitrogen-poor plants appear to allocate nitrogen to reproduction at the expense of vegetative organs throughout the life cycle.     

Interactive regulation of grass yield and chemical properties by defoliation,a salivary chemical,and inorganic nutrition

Summary Clones of 2 C₄ grass species, Sprobolus ioclados and S. pyramidalis, were obtained from more and less heavily grazed grasslands, respectively, in Tanzania's Serengeti National Park. Plants were grown in a factorial experiment to determine the effects of severe defoliation, nutrient limitation, and a salivary chemical (thiamine) on plant growth, nitrogen content, and non-structural carbohydrate content. The experimental design included: (1) species; (2) clipping, with plants either unclipped or clipped weekly to a height of 5 cm; (3) 0.2 ml of distilled water of 0.2 ml of 10 ppb aqueous thiamine sprayed on plants from an atomizer after clipping and identical treatments at the same time to unclipped plants; (4) phosphorus (P) at 0.2 or 1 mM; (5) nitrogen (N) at 3 or 15 mM. Clipping was the major variable affecting plant growth. Total and litter yields were reduced to half and residual plant yield was reduced to 30% of the values for unclipped plants. Clipping interacted strongly with other variables since they commonly had minor effects on clipped plants and major effects on unclipped plants. Exceptions to this generalization were generally due to better performance by S. ioclados under clipping. Compared to lower treatment levels, higher treatment levels promoted total yield of unclipped plants by 52% for N, 43% for thiamine, and 33% for P. In general, thiamine had greater effects than P but lesser effects than N. Thiamine promoted yield and modified the chemical balance of plants by promoting carbohydrate (CHO) concentrations and reducing N concentrations. N and P deficiencies promoted CHO accumulation. Clipping promoted the N of leaves and crowns and reduced the N levels in roots. Leaf blade water and N contents were positively correlated with very little scatter. The slope of the line was different for S. ioclados and S. pyramidalis. Leaf blade water and CHO contents were negatively related but there was more scatter and the species could not be distinguished. The species from more heavily grazed grasslands was conspicuously more sensitive to thiamine application. The results indicated that leaf treatment with thiamine, the only likely source of which in natural grasslands is saliva deposited by feeding herbivores, can have major effects on plant yield and metabolic balances at very low application levels. But under defoliation levels as severe as those imposed in this experiment, which reduced above ground plant biomass to a fourth of the level produced by unclipped plants, growth was so strongly limited by defoliation that neither thiamine nor inorganic nutrients affected plant yield residual from clipping. Therefore, whether chemicals such as thiamine that may be introduced onto grass foliage by grazing ungulates and other herbivores will influence the growth of grazed plants will depend upon the grazing intensity associated with the transfer.     

Plant-herbivore interactions: Examination of potential effects of bison saliva on regrowth of Bouteloua gracilis (H.B.K.) lag

Summary Laboratory experiments were performed to determine whether regrowth of blue grama was affected by potential growth-promoting substances in saliva of North American bison. We observed no statistically significant effects of foliar application of whole bison saliva on net photosynthesis (P_N), root respiration (R_R), allocation patterns of photosynthetically fixed ¹⁴C, or regrowth rates over a 10-day period following clipping to various heights. In a 10-week experiment, there were no significant effects of saliva on leaf, crown or root growth or tiller production in plants clipped to heights of 6, 4 or 2 cm above crowns. Similarly, nitrogen-stressed plants failed to show significant changes in growth rates or tillering in response to saliva over a 3-week period. Clipped blue grama plants did exhibit significant compensatory growth responses, including higher P_N rates from 3–10 days following clipping and allocation of a higher proportion of current photosynthate to synthesis of new leaf tissue with increasing severity of defoliation. Nevertheless, unclipped plants invariably outproduced clipped plants following defoliation.     

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。     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。     

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.     

Interactive effects of soil-dwelling ants, ant mounds and simulated grazing on local plant community composition

Interactions between aboveground vertebrate herbivores and subterranean yellow meadow ants (Lasius flavus) can drive plant community patterns in grassland ecosystems. Here, we study the relative importance of the presence of ants (L. flavus) and ant mounds under different simulated grazing regimes for biomass production and species composition in plant communities. We set up a greenhouse experiment using intact soil cores with their associated vegetation.We found that plant biomass production in the short term was affected by an interaction between simulated grazing (clipping) and ant mound presence. Clipping homogenized production on and off mounds, while in unclipped situations production was higher off than on mounds. During the experiment, these differences in unclipped situations disappeared, because production on unclipped mounds increased. Plant species richness was on average higher in clipped treatments and patterns did not change significantly over the experimental period. Plant community composition was mainly affected by clipping, which increased the cover of grazing-tolerant plant species. The actual presence of yellow meadow ants did not affect plant community composition and production.We conclude that the interaction between ant mounds and clipping determined plant community composition and biomass production, while the actual presence of ants themselves was not important. Moreover, clipping can overrule effects of ant mounds on biomass production. Only shortly after the cessation of clipping biomass production was affected by ant mound presence, suggesting that only under low intensity clipping ant mounds may become important determining plant production. Therefore, under low intensity grazing ant mounds may drive the formation of small-scale plant patches.     

盐碱条件下刈割干扰对羊草的氮素分配策略及补偿生长的影响

土地盐碱化和过度放牧是制约松嫩平原畜牧业发展的两大因素,羊草是松嫩平原上的优势种,被认为具有较强的耐牧及耐盐碱能力.本文通过田间原位试验,以叶面涂抹标记¹⁵N-尿素的方法,研究了不同盐碱条件下刈割干扰对羊草的氮素分配策略及补偿生长的影响.结果表明: 总体上叶面新吸收的氮60%以上保留在地上部分.与不施盐碱无刈割处理的对照相比,单纯的盐碱胁迫使新吸收的氮在细根中的分配率显著增加了5.1%;而盐碱胁迫下,中度刈割使叶面新吸收的氮在地上部分的分配率增加了11.6%,地上及总生物量发生超补偿生长,但是重度刈割使叶面新吸收的氮在茎基部的分配率显著增加了9.5%,地上、细根及总生物量均表现为欠补偿生长.上述结果表明盐碱胁迫下,中度刈割干扰时羊草采取积极的再生策略,促进其超补偿生长,但在重度刈割时羊草会采取增加氮素在茎基部存储的相对保守的氮素分配和生长策略.     

Reproductive allocation in hermaphrodite and female plants of Sidalcea oregana SSP. spicata (Malvaceae) using four currencies

Reproductive allocation was investigated in female and hermaphrodite plants of gynodioecious Sidalcea oregana ssp. spicata. Total reproductive investment and partitioning of that investment was documented at the level of whole plants in terms of four ecologically relevant currencies: biomass, nitrogen, phosphorus, and potassium. Nutrient augmentations in the field confirmed that nutrients were limiting plant vegetative growth and propensity to flower; thus the use of these nutrients as currency was appropriate. Once the effects of plant size were removed, the sex morphs allocated similar total amounts of biomass, nitrogen, phosphorus, and potassium to reproduction, but partitioned those differentially. For any given individual size, females allocated larger proportions of their reproductive resource budgets to seeds. Hermaphrodites' reproductive investment in pollen and flowers was allocated at the expense of allocation to seeds. These data are relevant to the evolution of gynodioecy from hermaphroditism and support the hypothesis that females reallocate resources not spent on pollen to seeds.     

Net photosynthesis,root respiration,and regrowth of Bouteloua gracilis following simulated grazing

Summary Net photosynthesis (P_N), root respiration (R_R), and regrowth of Bouteloua gracilis (H.B.K.) Lag. were examined in the laboratory over a 10-day period following clipping to a 4-cm height to simulate grazing by large herbivores. Net photosynthesis rates of tissue remaining immediately following defoliation were only about 40% as great as preclipping rates. Three days after clipping, P_N rates of defoliated plants had increased to values about 21% greater (per unit leaf area) than those of unclipped controls and remained at that level through Day 10. No statistically significant changes in R_R occurred following defoliation. Biomass of unclipped plants nearly doubled during the 10-day study period, while that of defoliated plants increased 67%. Over half the new growth of defoliated plants was allocated to new leaf blades and only 18% to new roots, while only 33% of the new growth of control plants was allocated to new leaf blades but 29% went to new roots. As a consequence of increased P_N rates and increased carbon allocation to synthesis of additional photosynthetic tissue following defoliation, net CO₂ uptake per plant increased from 9% to 80% of that of the controls from Day 0 through Day 10.     

No allocation trade-offs between flowering and sproutingin the lignotuberous,Mediterranean shrub Erica australis

Trade-offs between allocation to sexual or vegetative regeneration capacity are well established as a driving force in the life history patterns of plants in fire-prone environments. However, it is not known whether such trade-offs exist in plants which after aboveground removing disturbances, such as fire, may regenerate by sexual (seeding) or asexual (sprouting) mechanisms. We evaluated whether in the fire-recruiting resprouter Erica australis, which after fire can regenerate by seedling establishment or resprouting, a larger investment in flowers and seeds prior to being disturbed by clipping its aboveground parts would decrease subsequent sprouting, that is, its vegetative regeneration capacity. We analysed the relationships between flower and seed production and the ensuing production and growth of sprouts of six plants from thirteen different sites in central-western Spain. We found no significant relationships between measures of sexual reproductive effort and resprout production and growth 6 months after clipping the aboveground parts of the plants. No evidence of trade-offs between sexual and asexual efforts was found. Furthermore, no significant relationship was found between lignotuber total non-structural carbohydrates and sexual reproductive effort. In addition, 2 years after the disturbance, resprout biomass was positively and significantly correlated with sexual reproductive effort prior to the disturbance. This indicates that growth of resprouts was higher at the sites where plants made a greater reproductive effort. The sites that were more favourable to producing flowers and seeds could also be more favourable to resprouting.     

EFFECTS OF CLIPPING AND FOUR LEVELS OF NITROGEN ON THE GAS EXCHANGE,GROWTH, AND PRODUCTION OF TWO EAST AFRICAN GRAMINOIDS

Kyllinga nervosa and Themeda triandra plants were subjected to different clipping and nitrogen availability regimes. Following an extended period of growth under these conditions, total biomass, gas exchange and several morphological parameters were measured. Kyllinga nervosa showed compensatory growth to moderate levels of clipping whereas any clipping reduced the total biomass of T. triandra. Unclipped plants of either species were unable to respond to increased levels of nitrogen. Clipped plants responded in an ambiguous fashion, with increased allocation to offtake (material removed by clipping) in both species. Total biomass of K. nervosa was highest at 15 mM nitrogen levels which are equivalent to field levels. Both photosynthesis and respiration rates were unaffected by nitrogen treatments. Photosynthesis was significantly reduced by the most severe clipping regime of K. nervosa, but was unaffected by clipping of T. triandra.