几种针阔叶树种不同光照强度下叶绿素荧光特性的研究

测定研究了油松、新疆杨等7种针阔叶树种不同光照强度下最大量子效率(ΦP)和实际量子效率(PhiPSⅡ)等叶绿素荧光特性的变化规律,进而研究了树木对光照强度适应性。结果表明,ΦP和PhiPSⅡ随光强度增加呈下降趋势,二者的光进程变化呈正相关直线。运用方程获得的量子效率最大点、光进程变化转折点和量子效率最小点,可评价树木光合量子效率的大小和对强光的适应性。依PhiPSⅡ最大值对被试树种光合量子效率排序为沙棘>白榆>新疆杨>白腊>油松>白木千>侧柏;依ΦP光进程变化转折点和量子效率最小点对应的光强度评价被试树木对强光的适应能力排序为新疆杨>白榆>侧柏>白腊>油松>沙棘>白木千,结果与被试树种生态特征基本一致。     

Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

Nitrogen resorption from senescing leaves of three salt marsh plant species

Seasonalvariation in leaf nitrogen of mature green and senescent leaves and nitrogenresorption efficiency in three plants (Spartina maritima, Halimioneportulacoides and Arthrocnemum perenne) of aTagus estuary salt marsh are reported. Total nitrogen concentrations in greenand senescent leaves were higher during winter (December and March). Soilinorganic nitrogen availability showed an opposite pattern with higherconcentrations during summer (June and September) when total leaf biomass washigher. Nitrogen resorption efficiency ranged between 31 and 76% andH. portulacoides was the plant that better minimizednitrogen loss by this process. Nitrogen resorption occurred mainly from thesoluble protein pool, although other fractions must have been broken down duringthe resorption process. No significant seasonal variation in nitrogen resorptionefficiency and no relation to leaf total nitrogen or soil nitrogen availabilitywere found. This suggests that the efficiency of the resorption process is notdetermined by the plant nitrogen status nor by the availability of the nutrientin the soil. Nevertheless, resorption from senescing leaves may play animportant role in the nitrogen dynamics of salt marsh plants and reduce thenitrogen requirements for plant growth.     

Nutrient resorption response to fire and nitrogen addition in a semi-arid grassland

Fire and nitrogen (N) addition, both widely used grassland restoration strategies, strongly influence community composition and ecosystem functioning. However, little is known about their effects on plant nutrient resorption from senescing leaves, especially in semi-arid ecosystems. We evaluated the effects of fire, N addition (5.25 g N m⁻² yr⁻¹) and their potential interactions on nutrient resorption in five plant species in a semi-arid grassland in northern China. Foliar nutrient concentrations and resorption proficiencies and efficiencies varied substantially among species and functional groups. Fire increased green leaf N concentration ([N]g) and decreased N resorption proficiency (N RP), P resorption proficiency (P RP) and P resorption efficiency (P RE). N addition led to higher [N]g and lower N resorption, whereas it did not affect P related responses. There was no interaction between fire and N addition to affect all response variables except for green leaf P concentration ([P]g). These results suggest that fire and N addition can influence ecosystem nutrient cycling directly by changing resorption patterns and litter quality. Given the substantial interspecific variations in nutrient content and resorption and the potentially changing community composition, both fire and N addition may have indirect impacts on ecosystem nutrient cycling in this semi-arid grassland.     

Nutrient resorption patterns of plant functional groups in a tropical savanna: variation and functional significance

Green and senesced leaf nitrogen (N) and phosphorus (P) concentrations of different plant functional groups in savanna communities of Kruger National Park, South Africa were analyzed to determine if nutrient resorption was regulated by plant nutritional status and foliar N:P ratios. The N and P concentrations in green leaves and the N concentrations in senesced leaves differed significantly between the dominant plant functional groups in these savannas: fine-leaved trees, broad-leaved trees and grasses. However, all three functional groups reduced P to comparable and very low levels in senesced leaves, suggesting that P was tightly conserved in this tropical semi-arid savanna ecosystem. Across all functional groups, there was evidence for nutritional control of resorption in this system, with both N and P resorption efficiencies decreasing as green leaf nutrient concentrations increased. However, specific patterns of resorption and the functional relationships between nutrient concentrations in green and senesced leaves varied by nutrient and plant functional group. Functional relationships between N concentrations in green and senesced leaves were indistinguishable between the dominant groups, suggesting that variation in N resorption efficiency was largely the result of inter-life form differences in green leaf N concentrations. In contrast, observed differences in P resorption efficiencies between life forms appear to be the result of both differences in green leaf P concentrations as well as inherent differences between life forms in the fraction of green leaf P resorbed from senescing leaves. Our results indicate that foliar N:P ratios are poor predictors of resorption efficiency in this ecosystem, in contrast to N and P resorption proficiencies, which are more responsive to foliar N:P ratios.     

热带雨林三种树苗叶片光合机构对光强的适应

对生长在不同光强(自然日光的8％,25％,50％)下西双版纳热带雨林3种木本植物团花(Anthocephalus chinensis)、玉蕊(Barringtonia pendala)和藤黄(Garrcinia hanburyi)幼苗光合机构的研究表明,随着生长光强的升高,植物叶片的光饱和点、补偿点、净光合速率和非光化学淬灭系数(NPQ)升高,而表现量子效率(AQY)、有效光化学量子产量(Fv／Fm)、光化学淬灭系数(qP)下降．在抗氧化系统中,超氧化物歧化酶(SOD)、抗坏血酸过氧化酶(APX)活性随着光强的升高而升高,而过氢化物酶(CAT)活性与生长光强的变化不一致．抗坏血酸(AsA)含量随着光强的升高而急剧上升。最能反映PFD的变化．可以认为,除与叶黄素循环有关的热耗散增大之外,植物叶片抗氧化系统的加强也是响应强光的一种保护措施．     

4种阔叶树种叶中氮和磷的季节动态及其转移

从叶完全展开到生长季结束 ,对常绿阔叶树种日本米槠 (Castanopsis cuspidata(Thunb.) Schottky)和具柄冬青 (Ilexpedunculosa Miq)及落叶阔叶树种栎 (Quercus serrata Murr.)和栓皮栎 (Quercus variabilis Blume)叶片的 N和 P浓度、含量和养分转移进行了测定 .在生长期中日本米槠新叶的 N浓度在 5月为 36 .6 g/ kg,然后降到 15 .5和 17.5 g/ kg之间 ,其老叶的N浓度波动于 10 .4和 13.1g/ kg的范围内 ,而具柄冬青新叶的 N浓度从 2 7.3下降到 16 .0 g/ kg,接着上升到 18.3g/ kg,其老叶的 N浓度在 12 .0到 15 .5 g/ kg的范围内。栎和栓皮栎的叶 N浓度分别从 2 8.8下降到 18.1g/ kg和从 2 8.5下降到17.5 g/ kg。日本米槠新叶的 N含量从 1.5 4下降到 1.35 g/ m2 ,接着上升到 1.5 0 g/ m2 ,其老叶 N含量从 1.36下降到1.0 0 g/ m2 ,接着上升到 1.2 1g/ m2 ,而具柄冬青新叶的 N含量从 2 .2 5下降到 1.6 0 g/ m2 ,接着上升到 2 .2 0 g/ m2 ,其老叶的 N含量从 2 .13下降到 1.6 5 g/ m2。栎和栓皮栎的叶 N含量分别从 2 .10下降到 1.2 8g/ m2和从 2 .95下降到 2 .13g/ m2。日本米槠新叶的 P浓度由 3.39g/ kg降到 1.12和 1.15 g/ kg之间 ,其老叶的 P浓度变化于 0 .6 6和 0 .88g/ kg的范围内 ,而具柄冬青新叶的     

Convergent tapering of xylem conduits in different woody species

A recent theoretical model (the West, Brown and Enquist, WBE model) hypothesized that plants have evolved a network of xylem conduits with a tapered structure (narrower conduits distally) which should minimize the cost of water transport from roots to leaves. Specific measurements are required to test the model predictions. We sampled both angiosperms and gymnosperms (50 trees) growing in different environments with heights ranging from 0.5 to 44.4 m, measuring variations of the xylem-conduit diameter from tree top to stem base. In all trees measured, mean hydraulically weighted conduit diameters (Dh) at the tree top were narrower than those at the stem base. In actively growing trees, the longitudinal variation of Dh showed a degree of tapering in agreement with WBE predictions, while trees close to their maximum height showed slightly lower conduit tapering. Comparing different species, a very good correlation was observed between degree of xylem tapering and tree height (r2 = 0.88; P < 0.0001) independently of any other variable (age, site, altitude, etc.). As predicted by WBE, sampled trees seemed to converge towards similar xylem conduit tapering. However, trees approaching their maximum height had a nonoptimal tapering which appeared insufficient to compensate for the progressive increase in tree height.     

Responses to light changes in tropical deciduous woody seedlings with contrasting growth rates

We evaluated the responses in growth, biomass allocation, photosynthesis and stomatal conductance, to changes in light in woody seedlings from the tropical deciduous forest in Mexico, which shows a highly seasonal rain pattern. We studied ten species, which differed by 30-fold in relative growth rate (RGR). We analyzed plant growth in two contrasting light levels during 52 days and two transfers: from high to low (HL) and from low to high (LH) light intensity, and the respective controls in high (HH) and low (LL) light for another 52 days. The photosynthetic capacity (A _max) and stomatal conductance were measured at the day of the transfer between light conditions and at the end of the experiment. Species with high RGR showed the largest changes in RGR in response to contrasting light conditions (HH/LL ratio), and species with low RGR showed low responses. The fast-growing species were the most plastic, followed by species with intermediate growth rates, with the slow-growing species being the least plastic. Fast-growing species achieved higher maximum photosynthetic capacities (A _max) and stomatal conductance and higher response to light than slow-growing species. Species with high RGR showed a low RGR HH/LH ratio, suggesting a large response of L plants when transfered to H. The RGR of the species were associated with species specific leaf area and with the response in the leaf area, net assimilation rate and leaf weight ratio, suggesting the importance of the leaf area produced and the leaf characteristics rather than root:shoot ratio in determining RGR. Considering that seed germination is expected at the beginning of the rainy period, seedlings of most of the species will experience high-light conditions during its early growth. There are large annual variations in the time required for canopy closure (35–75 days). The influence of these variations may have different effect on the species studied. Species with intermediate growth rate and intermediate response to light changes were less affected by light reduction than fast-growing species. The intermediate-RGR species Caesalpiniaeriostachys is the most abundant and widely distributed species, perhaps this could be in part due to its ability to acclimate to both light increases and decreases. The fast-growing species studied here can be found in open sites in the forest and in areas cleared for pasture growth. These fast-growing species eventually reach the canopy, although this may require several canopy openings during their lives, which implies juvenile shade tolerance. In the tropical deciduous forest juvenile pioneer trees also benefit from the temporary high light available caused by the dry period during the rainy season. The slow-growing species Celaenodendronmexicanum forms small patches of monospecific forest; the adult trees are not completely deciduous, and they retain their old leaves for a long time period before shedding. Thus seedlings of this species may receive lower levels of light, in agreement with its shade tolerance and its lower response to light increases. Received: 14 April 1997 / Accepted: 29 July 1997     

Early root growth plasticity in seedlings of three Mediterranean woody species

Using a 141 F₂ population generated from maize inbred B64 × teosinte Zea nicaraguensis cross, quantitative trait loci (QTLs) controlling aerenchyma formation in roots under non-flooding drained soil conditions were identified. Seedlings of Z. nicaraguensis formed clear aerenchyma in the cortex of adventitious roots in non-flooding conditions, whereas the maize inbred line B64 did not. In the F₂ population, the capacity to develop aerenchyma exhibited wide and continuous variation, suggesting the trait was controlled by multiple genes. A linkage map was developed using 85 SSR markers, covering 1,224 cM across all ten chromosomes. Composite interval mapping analysis revealed that four QTLs for aerenchyma formation under non-flooding conditions were located to two regions of chromosome 1 (identified as Qaer1.02-3 and Qaer1.07), chromosome 5 (Qaer5.09) and chromosome 8 (Qaer8.06-7), and these explained 46.5% of the total phenotypic variance. The multiple interval mapping approach identified additional QTLs on chromosomes 1 (Qaer1.01) and 5 (Qaer5.01). Using these results, it may be possible to use SSR markers linked to aerenchyma formation in a marker assisted selection approach to introduce aerenchyma formation in drained soil conditions into maize for the eventual development of flooding tolerant maize hybrids.     

Low leaf N and P resorption contributes to nutrient limitation in two desert shrubs

Both water and nutrients are limiting in arid environments, and desert plants have adapted to these limitations through numerous developmental and physiological mechanisms. In the Mono Basin, California, USA, co-dominant Sarcobatus vermiculatus and Chrysothamnus nauseosus ssp. consimilis are differentially N and P limited. We hypothesized that low leaf N resorption contributes to N-limitation in Sarcobatus and that low leaf P resorption contributes to P-limitation in Chrysothamnus. As predicted, Sarcobatus resorbed proportionally 1.7-fold less N than Chrysothamnus, but reduced leaf P in senescent leaves to lower levels than Chrysothamnus (8.0–10.8-fold lower based on leaf area or mass, respectively), consistent with N, but not P limitations in Sarcobatus. Again, as predicted, Chrysothamnus resorbed proportionally 2.0-fold less P than Sarcobatus yet reduced leaf N in senescent leaves to lower levels than Sarcobatus (1.8–1.3-fold lower based on leaf area or mass, respectively), consistent with P, but not N limitations in Chrysothamnus. Leaf N and P pools were approximately 50% of aboveground pools in both species during the growing season, suggesting leaf resorption can contribute significantly to whole plant nutrient retention. This was consistent with changes in leaf N vs. P concentration as plants grew from seedlings to adults. Our results support the conclusion that N-limitation in Sarcobatus and P-limitation in Chrysothamnus are in part caused by physiological (or other) constraints that prevent more efficient resorption of N or P, respectively. For these species, differential nutrient resorption may be a key physiological component contributing to their coexistence in this saline, low resource habitat.     

Time-course of photosynthetic induction in four tropical woody species grown in contrasting irradiance habitats

We investigated the photosynthetic induction time-course in species of different ecological groups grown in contrasting forest irradiance environments, gap and understorey, exposed to different darkness times in order to verify the plant capacity to exploit irradiance heterogeneity. Photosynthetic induction was studied in leaves of Bauhinia forficata and Guazuma ulmifolia (early succession species, ES), and Esenbeckia leiocarpa and Hymenaea courbaril (late succession species, LS). T₅₀ and T₉₀ (time estimates to attain 50 and 90 % of maximum net photosynthetic rate, respectively) varied according to the time of previous exposure to darkness and growth irradiance. In both darkness times of 10 and 30 min, T₅₀ was lower in the LS-than ES-species. These results, jointly with significant higher induction state of the leaves after 10 min of darkness, suggest that the LS-species has a higher potential to sunfleck utilization compared to ES-species, both grown in the understorey. After 10 and 30 min of darkness the differences between ecological groups were not clearly detected in the gap for T₅₀ and T₉₀, indicating that eco-physiological characteristics of each ecological group did not influence the induction time of the species evaluated herein. Thus the capacity to show phenotypic plasticity is not exclusive to an ecological group, but it is rather a more intrinsic feature related to the differential capacity of individuals.     

Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species

Juvenile leaves in high-light environments commonly appear red as a result of anthocyanin pigments, which play a photoprotective role during light-sensitive ontogenetic stages. The loss of anthocyanin during leaf development presumably corresponds to a decreased need for photoprotection, as photosynthetic maturation allows leaves to utilize higher light intensities. However, the relationship between photosynthetic development and anthocyanin decline has yet to be quantitatively described. In this study, anthocyanin concentration was measured against photopigment content, lamina thickness, anatomical development, and photosynthetic CO(2) exchange in developing leaves of three deciduous tree species. In all species, anthocyanin disappearance corresponded with development of c. 50% mature photopigment concentrations, c. 80% lamina thickness, and differentiation of the mesophyll into palisade and spongy layers. Photosynthetic gas exchange correlated positively with leaf thickness and chlorophyll content, and negatively with anthocyanin concentration. Species with more rapid photosynthetic maturation lost anthocyanin earliest in development. Chlorophyll a/b ratios increased with leaf age, and were lower than those of acyanic species, consistent with a shading effect of anthocyanin. These results suggest that anthocyanin reassimilation is linked closely with chloroplast and whole-leaf developmental processes, supporting the idea that anthocyanins protect tissues until light processing and carbon fixation have matured to balance energy capture with utilization.     

Effect of vesicular-arbuscular mycorrhizae on seedling growth of four tree species from the tropical deciduous forest in Mexico

The influence of vesicular-arbuscular mycorrhizae on the growth of seedlings of Caesalpinia eriostachys, Cordia alliodora, Ipomoea wolcottiana and Pithecellobium mangense was investigated in a greenhouse experiment conducted at the Biological Station of Chamela on the Pacific coast of Mexico. Dry biomass production, relative growth rate, root/shoot ratio and mycorrhizal dependency were quantified for 75-day-old seedlings. With the exception of the pioneer species I. wolcottiana, mycorrhizal infection resulted in increases in biomass production, relative growth rate and leaf area. The root/shoot ratios attained for the species, however, did not show a consistent trend with infection. Nevertheless, all species had root/shoot ratios below 1 with infection and only one, Cordia alliodora, had a ratio greater than 1 without infection. The two late successional species from the mature part of the forest, Caesalpinia eriostachys and P. mangense, showed a larger mycorrhizal dependency than the two associated with disturbed environments.     

Diurnal variation and interrelations of ecophysiological parameters in three peatland woody species under different weather and soil moisture conditions

Summary The diurnal patterns of twig xylem water potential, net photosynthesis rate, water use efficiency of photosynthesis, and stomatal and mesophyll conductance to CO₂ in tamarack, black spruce and swamp birch growing in a natural peatland in central Alberta, Canada, were examined. The relationships of photosynthesis to other ccophysiological parameters were investigated. Data were collected on three days with different weather and soil moisture conditions in the 1988 growing season. Day 1 was clear and warm and the ground water table was 7 cm above the average peat surface. Day 2 was clear and hot. Day 3 was cloudy but warm. On day 2 and day 3, the water tables were in the normal range for that season. Major findings were: 1) Soil flooding depressed photosynthesis in tamarack and black spruce. 2) Swamp birch was better adapted to flooding than tamarack or black spruce. 3) The trees experienced water stress in the afternoons of the two days with lower water table. 4) Changes in photosynthesis of the three species were primarily affected by changes in mesophyll conductance (g_m) and the response of photosynthesis to changes in g_m was similar for all three species.     

Seasonal reversibility of acclimation to irradiance in leaves of common box (Buxus sempervirens L.) in a deciduous forest

Understorey shade plants are seasonally exposed to dramatic changes in light conditions in deciduous forests related with the dynamics of the overstorey leaf phenology. These transitions are commonly followed by changes in herb plant communities, but shade-tolerant evergreen species must be able to adapt to changing light conditions. In this work we checked the photoprotective responses of evergreen species to acclimate to the shady summer environment and reversibly de-acclimate to a more illuminated environment after leaf fall on deciduous overstoreys. For that purpose we have followed the process of light acclimation in leaves of common box (Buxus sempervirens) during the winter to spring transition, which decrease irradiance in the understorey, and conversely during the transition from summer to autumn. Four parameters indicative of the structure and degree of acclimation of the photosynthetic apparatus have been studied: chlorophyll a/b ratio which is supposed to be inversely proportional to the antenna size, α/β-carotene which increases in shade acclimated leaves and the pools of α-tocopherol and xanthophyll cycle pigments (VAZ) which are two of the main photoprotection mechanisms in plants. Among these parameters, chlorophyll a/b ratio and VAZ pool responded finely to changes in irradiance indicating that modifications in the light harvesting size and photoprotective capacity contribute to the continuous acclimation and de-acclimation of long-lived evergreen leaves.     

Susceptibility to photoinhibition of three deciduous broadleaf tree species with different successional traits raised under various light regimes

The susceptibility to photoinhibition of tree species from three different successional stages were examined using chlorophyll fluorescence and gas exchange techniques. The three deciduous broadleaf tree species were Betula platyphylla var. japonica, pioneer and early successional, Quercus mongolica, intermediate shade‐tolerant and mid‐successional, and Acer mono, shade‐tolerant and late successional. Tree seedlings were raised under three light regimes: full sunlight (open), 10% full sun, and 5% full sun. Susceptibility to photoinhibition was assessed on the basis of the recovery kinetics of the ratio of vaviable to maximum fluorescence (F_v/F_m) of detached leaf discs exposed to about 2000 μmol m^?1 s^?1 photon flux density (PFD) for 2 h under controlled conditions (25 to 28 °C, fully hydrated). Differences in susceptibility to photodamage among species were not significant in the open and 10% full sun treatments. But in 5% full sun, B. platyphylla sustained a significantly greater photodamage than other species, probably associated with having the lowest photosynthetic capacity indicated by light‐saturated photosynthetic rate (B. platyphylla, 9·87, 5·85 and 2·82; Q. mongolica, 8·05, 6·28 and 4·41; A. mono, 7·93, 6·11 and 5·08 μmol CO₂ m^?1 s^?1for open, 10% and 5% full sun, respectively). To simulate a gap formation and assess its complex effects including high temperature and water stress in addition to strong light on the susceptibility to photoinhibition, we examined photoinhibition in the field by means of monitoring ΔF/F′_m on the first day of transfer to natural daylight. Compared with ΔF/F′_m in AM, the lower ΔF/F′_m in PM responding to lower PFD following high PFD around noon indicated that photoinhibition occurred in plants grown in 10 and 5% full sun. The diurnal changes of ΔF/F′_m showed that Q. mongolica grown in 5% full sun was less susceptible to photoinhibition than A. mono although they showed little differences both in photosynthetic capacity in intact leaves and susceptibility to photoinhibition based on leaf disc measurements. These results suggest that shade‐grown Q. mongolica had a higher tolerance for additional stresses such as high temperature and water stress in the field, possibly due to their lower plasticity in leaf anatomy to low light environment.     

Changes in nitrogen resorption traits of six temperate grassland species along a multi-level N addition gradient

Nitrogen (N) resorption from senescing leaves is an important mechanism of N conservation for terrestrial plant species, but changes in N-resorption traits over wide-range and multi-level N addition gradients have not been well characterized. Here, a 3-year N addition experiment was conducted to determine the effects of N addition on N resorption of six temperate grassland species belonging to three different life-forms: Stipa krylovii Roshev. (grass), Cleistogenes squarrosa (T.) Keng (grass), Artemisia frigida Willd. (semishrub), Melissitus ruthenica C.W.Wang (semishrub and N-fixer), Potentilla acaulis L. (forb) and Allium bidentatum Fisch.ex Prokh. (forb). Generally, N concentrations in green leaves increased asymptotically for all species. N concentrations in senescent leaves for most species (5/6) also increased asymptotically, except that the N concentration in senescent leaves of A. bidentatum was independent of N addition. N-resorption efficiency decreased with increasing N addition level only for S. krylovii and A. frigida, while no clear responses were found for other species. These results suggest that long-term N fertilization increased N uptake and decreased N-resorption proficiency, but the effects on N-resorption efficiency were species-specific for different temperate grassland species in northern China. These inter-specific differences in N resorption may influence the positive feedback between species dominance and N availability and thus soil N cycling in the grassland ecosystem in this region.