Light field heterogeneity among tussock grasses: Theoretical considerations of light harvesting and seedling establishment in tussocks and uniform tiller distributions

Although the tussock growth form of caespitose graminoids is widespread, the effect of this growth form on light interception and carbon gain of tillers has received little attention. Daily incident photosynthetic photon flux density (PFD_inc) and carbon gain in monospecific stands of tussock grasses were compared with those of a hypothetical distribution with the equivalent tiller density per total ground area, but evenly distributed rather than clumped in tussocks. This was computed for two tussock grasses Pseudoroegneria spicata (Pursh) A. Löve (bluebunch wheatgrass) and Agropyron desertorum (Fisch, ex Link) Schult. (creasted wheatgrass) at different plant densities. Daily PFD_inc and net photosynthesis (A) were greater if tillers were distributed uniformly rather than clumped in tussocks, except when the density of tussocks was so great as to approach a uniform canopy. When tussock density per ground area was low, much of the difference between tussock and uniform tiller densities in PFD_inc and A was due to shading within the tussocks; up to 50–60% of the potential carbon gain was lost in A. desertorum due to shading within tussocks. In a matrix of tussocks, the light field for establishing seedlings was very heterogeneous; potential A ranged from 7 to 96% relative to an isolated seedling. The mean of daily PFD_inc and A for seedlings in a tussock stand were nearly identical to the values in corresponding stands of uniform tiller distributions. It is hypothesized that the loss of A resulting from clumping tillers into tussocks is offset by benefits of protecting sequestered belowground resources from invasion by seedlings of competitors.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary The annual replacement of tillers of Agropyron desertorum (Fisch. ex Link) Schult., a grazing-tolerant, Eurasian tussock grass, was examined in the field following cattle grazing. Heavy grazing before internode (culm) elongation seldom affected tiller replacement. Heavy grazing during or after internode elongation, which elevates apical meristems, increased overwinter mortality of fall-produced tillers and reduced the number and heights of these replacement tillers. Unexpectedly, tussocks grazed twice within the spring growing season tended to have lower overwinter tiller mortality, greater tiller replacement, and larger replacement tillers than tussocks grazed only once in late spring. These responses of twice-grazed tussocks, however, were still less than those of ungrazed tussocks or tussocks grazed moderately in early spring. The presence of ungrazed tillers on partially grazed tussoks did not increase the replacement of associated grazed tillers relative to tillers on uniformly grazed plants. This result indicates that resource sharing among tillers, if present, is short-lived or ecologically unimportant in this species. Although A. desertorum is considered grazing-tolerant, tiller replacement on heavily grazed tussocks, particularly those grazed during or after internode elongation when apical meristems were removed, was usually inadequate for tussock maintenance. These observations at the tiller (ramet) level of organization in individual tussocks (genet) may explain the often noted reduction in stand (population) longevity with consistent heavy grazing.     

Spatial arrangement of tiller replacement in Agropyron desertorum following grazing

Summary The spatial arrangement of tiller replacement was assessed on grazed and ungrazed tussocks of Agropyron desertorum (Fisch. ex Link) Schult. for three annual cycles. Frequency distributions of the number of replacement tillers per single progenitor were also determined. Tiller replacement was usually greater on the perimeter of tussocks than within the core, with or without grazing. Replacement was inversely related to grazing intensity, both on the perimeter and within the core of tussocks. Heights of replacement tillers on the perimeter or within the core seldom differed. Furthermore, grazing seldom affected the number of replacement tillers per progenitor. Greater tillering on the perimeter than within the core indicates that the tussocks were expanding. Apparently, grazing neither enhances tussock expansion and subsequent disintegration, nor does it necessarily lead to patches of tillers (multiple tillering per progenitor) within tussocks of A. desertorum.     

Morphological Constraints of Shoot Demography of a Clonal Plant: Extra- and Intravaginal Tillers of Festuca rubra

Abstract Festuca rubra forms tillers in two different ways: extravaginally and intravaginally. Demography of these two tiller types was observed in seventeen selected tussocks of Festuca rubra s.s. over four growing seasons. Extravaginal tillers were bigger at birth and on the average produced twice as many daughter tillers per tiller. In general, the natality and mortality of extravaginal tillers were less regular than that of intravaginal tillers. Overall tillering rate per tiller was correlated with the density of the surrounding vegetation; mortality, natality and tiller life span were not. High density of the surrounding vegetation did not result in increased formation of extravaginal tillers. The proportion of the extravaginal tillers was not correlated with the density of the F. rubra tussocks. There is no evidence for foraging by extravaginal tillers, but they do act as founders of small clusters of tillers.     

Rapid recovery of kohekohe (

Soil conditions, vegetation features and soil fauna were recorded in montane tall tussock grassland dominated by narrow- leaved snow tussock Chionochloa rigida ssp. rigida up to 30 months after a spring fire. Burning reduced the stature of tussocks and the size and density of tillers in the first growing season. After two growing seasons, tussock canopy development and tiller size remained below those found in the unburnt grassland nearby. New tillers and tussocks established following the prolific fire-induced flowering one year after burning. After the fire and sheep grazing, intertussock cover became progressively dominated by introduced grasses and herbs. While soil pH, moisture content, bulk density, surface litter and total nematodes showed significant treatment (burning) effects, these properties also showed significant year-to-year variation. The greatest increase in any nematode group was in Paratylenchus, a distinctive genus widespread in tussock grasslands and apparently responsive to environmental fluctuation and root development; its population was 100x and 29x greater in the burned area than in the control area 16 and 30 months after burning. Subject to detailed testing, populations of mites and collembola may provide relatively simple indicators of recovery of ecosystem function of such grasslands after burning.     

Evapotranspiration measurements in Eriophorum vaginatum tussock tundra in Alaska

Evapotranspiration was measured periodically from late June through mid-August 1978 in Eriophorum vaginiatum tussock tundra near Eagle Creek, Alaska. The average evapotranspiration rates from tussock and intertussock areas were 0.8 mm d⁻¹ and 1.3 mm d⁻¹, respectively. Potential evaporation was calculated according to the Penman equation using microclimate data collected at the same time actual evapotranspiration was measured. Actual evapotranspiration was 0.56 ± 0.06 ( X ± SE, N = 10) of the potential evaporation from moss and 0.43 ± 0.08 ( X ± SE, N = 10) of the potential evaporation from E. vaginatum tussocks. Seventy three percent of the variability in the ratio of actual evapotranspiration to potential evaporation from tussocks was accounted for by linear regression on net radiation. Eighty nine percent of the variability in the same ratio for mosses was explained by linear regression on the vapor pressure deficit of the air and net radiation.     

Kobresia tibetica tussocks facilitate plant species inside them and increase diversity and reproduction

Many plant species grow inside tussocks of some graminoids, but the underlying mechanisms remain unclear. We address whether some species occur and flower mainly inside tussocks so that species diversity and sexual reproduction are higher inside than outside tussocks, and whether relieving biological and physical stress is the mechanism associated with the facilitative process. In a heavily grazed grassland on the eastern Tibetan Plateau, where both physical (due to high altitude) and biological conditions (due to heavy grazing) are extremely harsh, we investigated vegetation in paired plots inside and outside 150 Kobresia tibetica tussocks and measured tussock basal area (=plot area). We also measured temperatures at soil surface, 5 and 10 cm depth and the number of animals (yaks, sheep and horses) grazing inside and outside tussocks. Sixty-seven percent of the species occurred and 42% flowered more frequently inside than outside tussocks, but none less frequently. Inside tussocks 78% species flowered, but outside tussocks only 31% did. Consequently, number of species, number of flowering species and number of inflorescences were all markedly larger inside than outside tussocks. Differences in number of species, number of flowering species and number of inflorescences inside and outside tussocks increased with increasing tussock basal area. Soil temperatures were lower inside than outside tussocks, but grazing intensity was much larger outside tussocks. Therefore, tussocks of K. tibetica facilitated the species inside them likely by grazing prevention, but not by increasing warmth. This study provides evidence that plant species colonizing tussocks of graminoids can be facilitated by the tussock species, and facilitation by grazing prevention may be one mechanism causing the coexistence of the species inside tussocks.     

Soil cellulase, chitinase, and protease activity in Eriophorum vaginatum tussock tundra at Eagle Summit, Alaska

Cellulase, chitinase, and protease activities were examined in Eriophorum vaginatum tussocks at Eagle Summit, Alaska. Changes in the activities were recorded for tussocks of differing relative plant composition. Highest total activity was found in tussocks with at least 50% (relative surface) cover by mycorrhizal colonized shrubs. Total enzyme activity declined in tussocks as the variety of plants decreased. The lowest enzyme activity was found in the moss dominated tussock. Changes in the nature of cellulose hydrolysis suggested that decomposition may be primarily regulated by litter composition in some cases and soil environment in others.     

Plant communities affect the species–area relationship on Carex sempervirens tussocks

The species–area relationship (SAR) has been extensively studied in a wide range of plant communities, but very few studies have directly addressed how plant communities affect the SAR and what are the underlying mechanisms. Many graminoids form distinct tussocks where many other plant species grow, but no study has investigated whether the SAR holds true for the vegetation on tussocks. In four plant communities on an abandoned subalpine pasture in the Swiss National Park, we made releves on 600 tussocks of Carex sempervirens and measured tussock basal area and other tussock traits. In all four communities, species richness on C. sempervirens tussocks was strongly positively related to tussock basal area (R²0.74), while other tussock traits explained very little (R²<0.04). Slope and intercept of the SAR on C. sempervirens tussocks differed significantly among the four communities. This was because plant communities affected richness in smaller tussocks (basal diameter <10 cm) but not that in large tussocks (basal diameter10 cm). We conclude that the SAR holds true for vegetation on C. sempervirens tussocks and changes with plant communities. Changes in the SAR on C. sempervirens tussocks are very likely because smaller tussocks are less independent of the plant communities than the larger ones, regarding disturbance or nutrients.     

Vegetative structure,microclimate, and leaf growth of a páramo tussock grass species,in undisturbed,burned and grazed conditions

In neotropical alpine grasslands (páramo), the natural tussock grass vegetation is extensively grazed and occasionally burned. The low productivity of the tussock grass seems to be the reason for the disappearance of this growth form in the most frequently intervened areas. The structure, microclimate and leaf elongation rates of new emerging leaves were studied for the dominant tussock grass species Calamagrostis effusa, at an undisturbed, a moderately grazed (7 year after fire) and a heavily grazed (3.5 years after fire) site. In absence of grazing and burning, the tussocks had a high standing crop (1.07±0.09 kg DW · m^-2) and leaf area per projected tussock cover (LAI: 9.6±1.4). Two thirds of the total mass was dead and more than half of the leaves were in horizontal position. The tussock growth form protects the meristems from severe climatic conditions. At midday, the temperature was higher at meristem level than in the rest of the tussock. At this level, photosynthetic irradiance (PI) was almost extinct at 2.9±0.74% of PI above the vegetation. The red/far red ratio (R/FR) was strongly decreased. Initial leaf elongation of new born leaves was 2.3 mm · day^-1, and constant during the year; estimated net annual production was 198±73.8 g m^-2. At the moderately grazed and the heavily grazed study sites, the tussocks were smaller, greener and more erect than those at the undisturbed site. More PI reached the meristems and R/FR was higher at the base of grazed tussocks. Leaf elongation rates were lower. Most of the litter disappeared during the fires. The lower elongation rate of leaves in the grazed areas might be a response to defoliation, resulting in increased tillering and a lack growth associated with poor temperature insulation and more UV-B damage.     

Multiclonal tussocks in the grass Achnatherum splendens (Trinius) Nevskia (Poaceae)

Clonal plant species often form genetically diverse populations, even when sexual reproduction in a population is rarely observed. Here we test whether the spatially discrete clusters of plants (tussocks of graminoids) formed within populations of some clonal species can likewise be multiclonal. We sampled leaves of ramets (shoots) within 20 tussocks of the grass Achnatherum splendens in the Otindag Sandland in Inner Mongolia, China, and genotyped the ramets using standard molecular protocols. The 20 tussocks were allocated to three classes: (i) small, circular, (ii) large, circular and (iii) large, irregular. Most tussocks (80%) were multiclonal and some contained at least eight different clones. Irregularly shaped tussocks contained twice as many clones as circular tussocks; neither size nor cover within a tussock affected number of clones per tussock, and the smaller clones in a tussock showed no tendency to occur on the edge or near the center of a tussock. These patterns seem more consistent with formation of multiclonal tussocks by coalescence than by colonization. Therefore, individual tussocks, especially large, irregular ones, cannot a priori be treated as genetic individuals without assessing their genetic information in, e.g., population demography, genetics and evolution studies.     

Seedling density and seedling survival in Alaskan cotton grass tussock tundra

Previous workers have suggested that reproduction by seedling establishment is less important in the maintenance of tundra vegetation than in temperate areas, but few quantitative data are available. Seedling densities were studied at 11 tundra sites in or near disturbed and undisturbed cotton grass tussock tundra in Alaska, and numerous seedlings were found. Examination of seedling age structures suggests that at least Ledum palustre and probably also Empetrum nigrum do replace themselves by seed in undisturbed tundra. Seedling densities vary greatly depending on the substrate available for establishment.     

人工恢复与自然恢复模式下苔草草丘生态特征比较

本研究以哈尔滨太阳岛草丘湿地为对象,对比了人工恢复与自然恢复下苔草草丘个体和种群的生态特征,并分析其与环境因子的关系.结果表明:苔草植株生长随时间呈现先增加后降低的变化趋势(5—8月),初期(5—6月)生长迅速,6月达到峰值.人工恢复和自然恢复模式下,苔草草丘个体和种群特征差异显著:自然恢复下苔草叶面积、叶宽、单株鲜重、单株干重、丘墩高度、直径、丘顶面积、丘墩表面积、丘墩体积等苔草草丘个体特征均显著高于人工恢复,人工恢复下苔草草丘密度、盖度、生物量等种群特征显著高于自然恢复,物种多样性无显著差异.土壤含水量、水深、草丘密度、丘间距离是导致2种恢复模式下苔草草丘生长差异的主要因素,自然恢复区土壤含水量、水深、间距均显著高于人工恢复区,对草丘个体的形成和发育具有促进作用,人工恢复区高移栽密度是导致草丘密度、盖度、生物量高于自然恢复区的主要因素.建议未来开展苔草草丘湿地恢复和保护时,应参考自然恢复湿地中草丘的分布特点,适当调整丘间距离(54.22～117.89 cm)和种群密度(1.9～3.1 墩·m^-2),同时采取干旱区春季适当补水措施,保持适宜的土壤含水量和水深,促进苔草草丘的生长发育和快速恢复,维持其种群长期健康稳定.     

Age-states,population structure,and seedling regeneration of Chionochloa pallens in Canterbury alpine grasslands,New Zealand

Abstract. Age-states, population structure, and seedling regeneration of the grass Chionochloa pallens were investigated in eight alpine grassland stands representing the range of canopy cover (ca. 5–75 %) and basal area (112 - 3889 m²/ha^-1) in the headwaters of the Avoca catchment, Canterbury, New Zealand. In open-canopy stands (< 50 % canopy cover, < 2000 m²/ha^-1 basal area) C. pallens individuals were clearly distinguishable. Rela-tionshipsbetweenbasal diameter, number of tillers, height, and % crown death suggested four putative age-states: ‘seedlings‘, ‘juvenile tussocks’, ‘mature tussocks’ and ‘senescent tussocks’. Four types of population structure were identified depending onthe proportions of seedlings and juvenile and mature tussocks. C. pallens seedlings were present in seven stands at densities ranging from 0.6 to 123.4 / m². They were most frequent on microsites protected from frost-heave by short herbaceous vegetation or litter, but with little competition from taller vegetation. Differences in seedling densities between stands partly reflected the proximity and abundance of seed sources and degree of protection from frost-heave. C. pallens population structures appear to be influenced by the frequency and magnitude of geomorphic disturbances and browsing by introduced animals.     

Antarctic moss carpets facilitate growth of Deschampsia antarctica but not its survival

The vegetation of the Antarctic tundra is dominated by mosses and lichens. Deschampsia antarctica, the Antarctic hairgrass, is one of two vascular plant species which grow along the west coast of the Antarctic Peninsula. However, little is known about its recruitment and interaction with non-vascular tundra plants. Although several authors propose that tolerance and/or competition should be the main forms of interaction between moss carpets and D. antarctica, no relevant studies exist so far. We investigated whether positive interactions are predominant at the Shetland Islands and the west coast of the Antarctic Peninsula and focussed on the role that moss carpets play in the recruitment of D. antarctica. Across the studied zone, D. antarctica showed a significant association with moss carpets, with higher frequencies as well as more and larger individuals than on bare ground. At one site, we conducted moss removal and seedlings transplant experiments to assess the relevance of the moss carpets for different life stages of hairgrass. All experimental individuals survived until the following summer whether the moss carpet was removed or not, but growth rate was significantly lower in tussocks with moss carpets removed. Likewise, tiller size was higher in plants growing in moss carpets than on bare ground. The detected positive interactions with mosses seem to be important for the expansion of D. antarctica, raising the question about their importance under future climate change scenarios.     

Microdistribution and water loss resistances of selected bryophytes in an Alaskan Eriophorum tussock tundra

Bryophyte species distributions were analyzed with respect to microtopography at an alpine tundra site in central Alaska which is dominated by tussocks of Eriophorum vaginatum . Bryophyte distributions were found to be significantly correlated with slope but not with azimuth. Different types of tussocks and hollows and mats between tussocks also supported different bryophyte floras. Water loss resistances of three species of moss did not account for differences in their distributions.     

Leaf and Tiller Production of Prairie Grass (Bromus willdenowii Kunth) and Two Ryegrass (Lolium) Species

A detailed morphological study of three prairie grass cultivars(Bromus willdenowii Kunth) was conducted under ‘vegetative’and ‘reproductive’ growth conditions (short andlong photoperiods) and at different temperatures. Perennialryegrass (Lolium perenne L.) and Westerwolds ryegrass (Loliummuhiflorum Lam.) were compared during vegetative growth. Prairie grass had higher leaf appearance rates (leaves per tillerper day) and lower site filling (tillers per tiller per leafappearance interval) than the ryegrass species. Tillering rates(tillers per tiller per day) were also lower, except under vegetativeconditions at 4C. Low tiller number in prairie grass was notdue to lack of tiller sites but a result of poor filling ofthese sites. Lower site filling occurred because of increaseddelays in appearance of the youngest axillary tiller and lackof axillary tillers emerging from basal tiller buds. In prairiegrass, no tillers came from coleoptile buds while only occasionallydid prophyll buds develop tillers. Low tiller number in prairiegrass was compensated for by greater tiller weight. Prairiegrass had more live leaves per tiller, greater area per leafand a high leaf area per plant. Considerable variation between cultivars was found in prairiegrass. The cultivar ‘Bellegarde’ had high leaf appearance,large leaves and rapid reproductive development, but had lowlevels of site filling, tillering rate, final tiller numberand herbage quality during reproductive growth. ‘Primabel’tended to have the opposite levels for these parameters, while‘Grasslands Matua’ was intermediate and possiblyprovided the best balance of all plant parameters. prairie grass, Bromus willdenowii Kunth, perennial ryegrass, Lolium perenne L., Westerwolds ryegrass, Lolium multiflorum Lam., temperature, photoperiod, leaf appearance, leaf area, tillering, site filling, tillering sites, yield     

Reproductive effort in cotton grass tussock tundra

Eriophorum vaginatum ssp. spissum is a dominant plant species of undisturbed cotton grass tussock tundra in Alaska. It also quickly invades and dominates recently disturbed sites. The hypothesis tested in this research was that the success of E. vaginatum on disturbed sites might be achieved through a higher allocation of biomass to reproductive structures relative to other tundra species. Reproductive allocation of tundra plants in general also was compared with plants of the temperate zone. The results indicate that E. vaginatum is about average among the common tundra species in terms of total reproductive allocation, allocation to seeds, and the proportion of total reproductive allocation that is accounted for by viable seeds. Tundra species, on a relative basis, allocate less biomass to all reproductive structures than temperate species but not necessarily less biomass to the output of viable seeds.     

Tillering in grain sorghum over a wiide range of population densities: modelling dynamics of tiller fertility

The prediction of tillering is poor or absent in existing sorghum crop models even though fertile tillers contribute significantly to grain yield. The objective of this study was to identify general quantitative relationships underpinning tiller dynamics of sorghum for a broad range of assimilate availabilities. Emergence, phenology, leaf area development and fertility of individual main culms and tillers were quantified weekly in plants grown at one of four plant densities ranging from two to 16 plants m(-2). On any given day, a tiller was considered potentially fertile (a posteriori) if its number of leaves continued to increase thereafter. The dynamics of potentially fertile tiller number per plant varied greatly with plant density, but could generally be described by three determinants, stable across plant densities: tiller emergence rate aligned with leaf ligule appearance rate; cessation of tiller emergence occurred at a stable leaf area index; and rate of decrease in potentially fertile tillers was linearly related to the ratio of realized to potential leaf area growth. Realized leaf area growth is the measured increase in leaf area, whereas potential leaf area growth is the estimated increase in leaf area if all potentially fertile tillers were to continue to develop. Procedures to predict this ratio, by estimating realized leaf area per plant from intercepted radiation and potential leaf area per plant from the number and type of developing axes, are presented. While it is suitable for modelling tiller dynamics in grain sorghum, this general framework needs to be validated by testing it in different environments and for other cultivars.     

Tiller population dynamics of reciprocally transplanted Eriophorum vaginatum L. ecotypes in a changing climate

Moist tussock tundra, dominated by the sedge Eriophorum vaginatum L., covers approximately 3.36 × 10⁸ km² of arctic surface area along with large amounts of subarctic land area. Eriophorum vaginatum exhibits ecotypic differentiation along latitudinal gradients in Alaska. While ecotypic differentiation may be beneficial during periods of climate stability, it may be detrimental as climate changes, causing adaptational lag. Following harvest of a 30-year reciprocal transplant experiment, age-specific demographic data on E. vaginatum tillers were collected to parameterize a Leslie matrix. Yellow Taxi analysis, based on Tukey's Jackknife, was used to determine mean pseudovalues of tiller population growth rate () for four source populations of E. vaginatum tussocks that were transplanted to each of three gardens along a latitudinal gradient. Source populations responded differentially along the latitudinal gradient. Survival and daughter tiller production influenced differences seen at the mid-latitude garden, and the overall tiller population performance was generally improved by northward transplanting relative to southward transplanting. A comparison of home-source and away-source within the same transplant garden indicates no home-site advantage. Although populations were still growing when transplanted to home-sites ( = 1.056), tiller population growth rate increased as ΔGDD became more negative relative to the home site (i.e., as tussocks were transplanted north). These results imply that populations are affected by climate gradients in a manner consistent with adaptational lag. This study documenting the response of high-latitude ecotypes to climate gradients may be an indication of the possible future effects of climate shift in more southern latitudes.