Plants Mediate the Sensitivity of Soil Respiration to Rainfall Variability

Soil respiration from grasslands plays a critical role in determining carbon dioxide (CO₂) feedbacks between soils and the atmosphere. In these often mesic systems, soil moisture and temperature tend to co-regulate soil respiration. Increasing variance of rainfall patterns may alter aboveground–belowground interactions and have important implications for the sensitivity of soil respiration to fluctuations in moisture and temperature. We conducted a set of field experiments to evaluate the independent and interactive effects of rainfall variability and plant–soil processes on respiration dynamics. Plant removal had strong effects on grassland soils, which included altered CO₂ flux owing to absence of root respiration; increased soil moisture and temperature; and reduced availability of dissolved organic carbon (DOC) for heterotrophic respiration by microorganisms. These plant-mediated effects interacted with our rainfall variability treatments to determine the sensitivity of soil respiration to both moisture and temperature. Using time-series multiple regression, we found that plants dampened the sensitivity of respiration to moisture under high variability rainfall treatments, which may reflect the relative stability of root contributions to total soil respiration. In contrast, plants increased the sensitivity of respiration to temperature under low variability rainfall treatment suggesting that the environmental controls on soil CO₂ dynamics in mesic habitats may be context dependent. Our results provide insight into the aboveground–belowground mechanisms controlling respiration in grasslands under variable rainfall regimes, which may be important for predicting CO₂ dynamics under current and future climate scenarios.     

Altering Rainfall Timing and Quantity in a Mesic Grassland Ecosystem: Design and Performance of Rainfall Manipulation Shelters

Global climate change is predicted to alter growing season rainfall patterns, potentially reducing total amounts of growing season precipitation and redistributing rainfall into fewer but larger individual events. Such changes may affect numerous soil, plant, and ecosystem properties in grasslands and ultimately impact their productivity and biological diversity. Rainout shelters are useful tools for experimental manipulations of rainfall patterns, and permanent fixed-location shelters were established in 1997 to conduct the Rainfall Manipulation Plot study in a mesic tallgrass prairie ecosystem in northeastern Kansas. Twelve 9 x 14–m fixed-location rainfall manipulation shelters were constructed to impose factorial combinations of 30% reduced rainfall quantity and 50% greater interrainfall dry periods on 6 x 6–m plots, to examine how altered rainfall regimes may affect plant species composition, nutrient cycling, and above- and belowground plant growth dynamics. The shelters provided complete control of growing season rainfall patterns, whereas effects on photosynthetic photon flux density, nighttime net radiation, and soil temperature generally were comparable to other similar shelter designs. Soil and plant responses to the first growing season of rainfall manipulations (1998) suggested that the interval between rainfall events may be a primary driver in grassland ecosystem responses to altered rainfall patterns. Aboveground net primary productivity, soil CO₂ flux, and flowering duration were reduced by the increased interrainfall intervals and were mostly unaffected by reduced rainfall quantity. The timing of rainfall events and resulting temporal patterns of soil moisture relative to critical times for microbial activity, biomass accumulation, plant life histories, and other ecological properties may regulate longer-term responses to altered rainfall patterns.     

Productivity responses to altered rainfall patterns in a C<Subscript>4</Subscript>-dominated grassland

Rainfall variability is a key driver of ecosystem structure and function in grasslands worldwide. Changes in rainfall patterns predicted by global climate models for the central United States are expected to cause lower and increasingly variable soil water availability, which may impact net primary production and plant species composition in native Great Plains grasslands. We experimentally altered the timing and quantity of growing season rainfall inputs by lengthening inter-rainfall dry intervals by 50%, reducing rainfall quantities by 30%, or both, compared to the ambient rainfall regime in a native tallgrass prairie ecosystem in northeastern Kansas. Over three growing seasons, increased rainfall variability caused by altered rainfall timing with no change in total rainfall quantity led to lower and more variable soil water content (0–30 cm depth), a ~10% reduction in aboveground net primary productivity (ANPP), increased root to shoot ratios, and greater canopy photon flux density at 30 cm above the soil surface. Lower total ANPP primarily resulted from reduced growth, biomass and flowering of subdominant warm-season C₄ grasses while productivity of the dominant C₄ grass Andropogon gerardii was relatively unresponsive. In general, vegetation responses to increased soil water content variability were at least equal to those caused by imposing a 30% reduction in rainfall quantity without altering the timing of rainfall inputs. Reduced ANPP most likely resulted from direct effects of soil moisture deficits on root activity, plant water status, and photosynthesis. Altered rainfall regimes are likely to be an important element of climate change scenarios in this grassland, and the nature of interactions with other climate change elements remains a significant challenge for predicting ecosystem responses to climate change.     

Belowground net primary productivity and biomass allocation of a grassland in Inner Mongolia is affected by grazing intensity

The root system of permanent grasslands is of outstanding importance for resource acquisition. Particularly under semi-arid conditions, the acquisition of water and nutrients is highly variable during the vegetation growth period and between years. Additionally, grazing is repeatedly disturbing the functional equilibrium between the root system and the transpiring leaf canopy. However, very few data is available considering grazing effects on belowground net primary productivity (BNPP) and root-shoot dry mass allocation in natural grassland systems. We hypothesise that grazing significantly reduces BNPP due to carbon reallocation to shoot growth. Root biomass and BNPP were estimated by soil coring in 2004, 2005 and 2006 and from ingrowth cores in 2005 and 2006 at one site which has been protected from grazing since 1979 (UG79), at one winter grazing (WG), and one heavily grazed (HG) site. BNPP was estimated from the summation of significant increments of total and live root biomass and from accumulated root biomass of ingrowth cores. Belowground biomass varied from 1,490–2,670 g m^?2 and was significantly lower under heavy grazing than at site UG79. Root turnover varied from 0.23 to 0.33 year^?1 and was not significantly different between sites. Heavy grazing significantly decreased live root biomass and BNPP compared to site UG79. Taking BNPP estimates from live root biomass dynamics and ingrowth cores as the most reliable values, the portion of dry mass allocated belowground relative to total net primary productivity (BNPP/NPP) varied between 0.50–0.66 and was reduced under heavy grazing in 2005, but not in 2006. The positive correlation between cumulative root length density of ingrowth cores and leaf dry matter suggests that the ingrowth core method is suitable for studying BNPP in this semi-arid steppe system. Grazing effects on BNPP and BNPP/NPP should be considered in regional carbon models and estimates of belowground nutrient cycling.     

IMPORTANCE OF TIME OF GERMINATION AND SOIL DEPTH ON GROWTH OF PROSOPIS GLANDULOSA (LEGUMINOSAE) SEEDLINGS IN THE PRESENCE OF A C4 GRASS

In an investigation of the causes of the invasion of woody plants into grasslands, competition between seedlings of Prosopis glandulosa Torr. and Bouteloua curtipendula (Michx.) Torr. was examined. Introduction of P. glandulosa into a B. curtipendula neighborhood significantly reduced P. glandulosa dry mass when compared to P. glandulosa growth alone. The greater the time interval from P. glandulosa germination to addition of B. curtipendula, the less interference the grass had on woody plant growth. Reciprocally, the greater the time interval from B. curtipendula germination to addition of P. glandulosa, the more interference the grass had on woody plant growth. Prosopis glandulosa belowground dry mass was <0.02 g (all in the upper 30 cm of soil) when planted after B. curtipendula at any soil depth, but if planted alone its root dry mass ranged from 2 to 8 g depending on depth. Prosopis glandulosa seedling dry mass increased linearly with soil depth, while B. curtipendula dry mass reached a plateau. In general, belowground dry mass of P. glandulosa was distributed throughout the soil depth examined (decreasing with depth), while 80% of B. curtipendula dry mass was found in the upper 30 cm of soil, suggesting a partitioning of soil resources. Data suggest that P. glandulosa and perhaps other shade-intolerant woody species that establish in grasslands do so in disturbances or vegetation gaps. Gaps may close, but by this time woody plant roots are below grass roots, thus partitioning soil resources and reducing interspecific competition.     

Interactive Effects of Time, CO2, N, and Diversity on Total Belowground Carbon Allocation and Ecosystem Carbon Storage in a Grassland Community

Predicting if ecosystems will mitigate or exacerbate rising CO₂ requires understanding how elevated CO₂ will interact with coincident changes in diversity and nitrogen (N) availability to affect ecosystem carbon (C) storage. Yet achieving such understanding has been hampered by the difficulty of quantifying belowground C pools and fluxes. Thus, we used mass balance calculations to quantify the effects of diversity, CO₂, and N on both the total amount of C allocated belowground by plants (total belowground C allocation, TBCA) and ecosystem C storage in a periodically burned, 8-year Minnesota grassland biodiversity, CO₂, and N experiment (BioCON). Annual TBCA increased in response to elevated CO₂, enriched N, and increasing diversity. TBCA was positively related to standing root biomass. After removing the influence of root biomass, the effect of elevated CO₂ remained positive, suggesting additional drivers of TBCA apart from those that maintain high root biomass. Removing root biomass effects resulted in the effects of N and diversity becoming neutral or negative (depending on year), suggesting that the positive effects of diversity and N on TBCA were related to treatment-driven differences in root biomass. Greater litter production in high diversity, elevated CO₂, and enhanced N treatments increased annual ecosystem C loss in fire years and C gain in non-fire years, resulting in overall neutral C storage rates. Our results suggest that frequently burned grasslands are unlikely to exhibit enhanced C sequestration with increasing atmospheric CO₂ levels or N deposition.     

The relationship between outbreaks, rainfall and low density populations of the African armyworm, Spodoptera exempta, in Kenya

This study has demonstrated that low numbers of African armyworms (Spodoptera exempta) (Walker) (Lepidoptera: Noctuidae) can be found throughout the year in parts of the coastal and highland regions of Kenya where there is frequent rainfall. During the study period the numbers of moths caught in pheromone traps in these regions built up during the short and long rains, and decreased dramatically during the intervening dry seasons. There was a lag of one to two months between the peaks for rainfall and moth numbers. This contrasts with the situation at sites of the seasonal outbreaks of armyworm larvae, where a sudden preceding influx of moths coincides with the rainfall. A positive correlation was found, for the long rains seasons only, between the number of armyworm outbreaks throughout the country and the peak numbers of moths in coastal and highland regions.It is suggested therefore that the peak numbers of moths trapped in the eastern highlands and coastal regions during the rainy seasons arise principally from outbreaks in the extensive seasonal grasslands. These grassland areas diminish considerably during the prolonged dry periods between the two rainy seasons, and it is only the eastern highland and coastal regions that are likely to provide suitable habitats for breeding during the dry seasons. The significance of these populations for initiating the first outbreaks of the following season is discussed.     

Contrasting above‐ and belowground sensitivity of three Great Plains grasslands to altered rainfall regimes

Intensification of the global hydrological cycle with atmospheric warming is expected to increase interannual variation in precipitation amount and the frequency of extreme precipitation events. Although studies in grasslands have shown sensitivity of aboveground net primary productivity (ANPP) to both precipitation amount and event size, we lack equivalent knowledge for responses of belowground net primary productivity (BNPP) and NPP. We conducted a 2‐year experiment in three US Great Plains grasslands – the C₄‐dominated shortgrass prairie (SGP; low ANPP) and tallgrass prairie (TGP; high ANPP), and the C₃‐dominated northern mixed grass prairie (NMP; intermediate ANPP) – to test three predictions: (i) both ANPP and BNPP responses to increased precipitation amount would vary inversely with mean annual precipitation (MAP) and site productivity; (ii) increased numbers of extreme rainfall events during high‐rainfall years would affect high and low MAP sites differently; and (iii) responses belowground would mirror those aboveground. We increased growing season precipitation by as much as 50% by augmenting natural rainfall via (i) many (11–13) small or (ii) fewer (3–5) large watering events, with the latter coinciding with naturally occurring large storms. Both ANPP and BNPP increased with water addition in the two C₄ grasslands, with greater ANPP sensitivity in TGP, but greater BNPP and NPP sensitivity in SGP. ANPP and BNPP did not respond to any rainfall manipulations in the C₃‐dominated NMP. Consistent with previous studies, fewer larger (extreme) rainfall events increased ANPP relative to many small events in SGP, but event size had no effect in TGP. Neither system responded consistently above‐ and belowground to event size; consequently, total NPP was insensitive to event size. The diversity of responses observed in these three grassland types underscores the challenge of predicting responses relevant to C cycling to forecast changes in precipitation regimes even within relatively homogeneous biomes such as grasslands.     

Limiting processes for perennial plant reintroduction to restore dry grasslands

In restored grasslands of southern Europe, perennial plants remain highly underrepresented compared with the reference ecosystems. We tested various treatments to reintroduce common perennial plant species (Brachypodium retusum, Poaceae, and Thymus vulgaris, Lamiaceae), which are usually not or poorly reintroduced via soil and hay transfer. Treatments included microenvironmental manipulations (rock cover and plant interactions) and two grazing intensities. Target perennial species were transplanted in 2002 in the reference grassland ecosystem (intact grassland area used as a control) and in two abandoned fields. Survival was assessed in June 2003 and June 2004. Target species shoot and root biomass were measured in June 2004. Grazing greatly reduced the survival and biomass of both target species and its effects were reinforced by summer drought: plants that did not establish well enough during the autumn and spring did not survive summer. The restored rock cover had a mild positive effect, particularly on B. retusum. There were no negative or positive plant neighbor interactions in the steppe, while there was competition in both abandoned fields. Competition was particularly intense in the abandoned melon field, composed of a dense sward of annual grasses (Bromus sp.). In order to reintroduce perennial species to dry grasslands, the ideal combination of treatments is to exclude or reduce grazing during the first year to allow seedlings to establish and to recreate adequate microenvironmental conditions. Reducing competition from arable weeds may help but is not essential in such dry grasslands.     

Differential effect of drought regimes on the seedling performance of six floodplain grassland species

• The performance of seedlings is crucial for the survival and persistence of plant populations. Although drought frequently occurs in floodplains and can cause seedling mortality, studies on the effects of drought on seedlings of floodplain grasslands are scarce. We tested the hypotheses that drought reduces aboveground biomass, total biomass, plant height, number of leaves, leaf area and specific leaf area (SLA), and increases root biomass and root‐mass fraction (RMF) and that seedlings from species of wet floodplain grasslands are more affected by drought than species of dry grasslands.
• In a greenhouse study, we exposed seedlings of three confamilial pairs of species (Pimpinella saxifraga, Selinum carvifolia, Veronica teucrium, Veronica maritima, Sanguisorba minor, Sanguisorba officinalis) to increasing drought treatments. Within each plant family, one species is characteristic of wet and one of dry floodplain grasslands, confamilial in order to avoid phylogenetic bias of the results.
• In accordance with our hypotheses, drought conditions reduced aboveground biomass, total biomass, plant height, number of leaves and leaf area. Contrary to our hypotheses, drought conditions increased SLA and decreased root biomass and RMF of seedlings. Beyond the effects of the families, the results were species‐specific (V. maritima being the most sensitive species) and habitat‐specific. Species indicative of wet floodplain grasslands appear to be more sensitive to drought than species indicative of dry grasslands.
• Because of species‐ and habitat‐specific responses to reduced water availability, future drought periods due to climate change may severely affect some species from dry and wet habitats, while others may be unaffected.

     

青藏高原高寒灌丛生态系统草本层生物量分配格局

青藏高原高寒灌丛生态系统生物量分配的研究相对较少,尤其是其草本层。为了探究高寒灌丛生态系统草本层生物量分配特征及其影响因素,分析了青藏高原东北部灌丛生态系统的49个高寒灌丛样地的草本层地上与地下生物量特征及其气候因子之间的关系。结果表明1)草本层地上生物量与地下生物量分别为121.1,342.8 g/m2均大于高寒草地的地上生物量与地下生物量。2)草本层的根冠比为3.6低于高寒草地的根冠比。3)地上生物量与地下生物量之间呈现幂函数的关系y=8.0x0.83(R2=0.48,P0.001)。4)根冠比与年均温度、年均降雨量之间没有显著的相关关系。     

Increased rainfall variability and reduced rainfall amount decreases soil CO2 flux in a grassland ecosystem

Predicted climate changes in the US Central Plains include altered precipitation regimes with increased occurrence of growing season droughts and higher frequencies of extreme rainfall events. Changes in the amounts and timing of rainfall events will likely affect ecosystem processes, including those that control C cycling and storage. Soil carbon dioxide (CO₂) flux is an important component of C cycling in terrestrial ecosystems, and is strongly influenced by climate. While many studies have assessed the influence of soil water content on soil CO₂ flux, few have included experimental manipulation of rainfall amounts in intact ecosystems, and we know of no studies that have explicitly addressed the influence of the timing of rainfall events. In order to determine the responses of soil CO₂ flux to altered rainfall timing and amounts, we manipulated rainfall inputs to plots of native tallgrass prairie (Konza Prairie, Kansas, USA) over four growing seasons (1998–2001). Specifically, we altered the amounts and/or timing of growing season rainfall in a factorial combination that included two levels of rainfall amount (100% or 70% of naturally occurring rainfall quantity) and two temporal patterns of rain events (ambient timing or a 50% increase in length of dry intervals between events). The size of individual rain events in the altered timing treatment was adjusted so that the quantity of total growing season rainfall in the ambient and altered timing treatments was the same (i.e. fewer, but larger rainfall events characterized the altered timing treatment). Seasonal mean soil CO₂ flux decreased by 8% under reduced rainfall amounts, by 13% under altered rainfall timing, and by 20% when both were combined (P<0.01). These changes in soil CO₂ flux were consistent with observed changes in plant productivity, which was also reduced by both reduced rainfall quantity and altered rainfall timing. Soil CO₂ flux was related to both soil temperature and soil water content in regression analyses; together they explained as much as 64% of the variability in CO₂ flux across dates under ambient rainfall timing, but only 38–48% of the variability under altered rainfall timing, suggesting that other factors (e.g. substrate availability, plant or microbial stress) may limit CO₂ flux under a climate regime that includes fewer, larger rainfall events. An analysis of the temperature sensitivity of soil CO₂ flux indicated that temperature had a reduced effect (lower correlation and lower Q₁₀ values) under the reduced quantity and altered timing treatments. Recognition that changes in the timing of rainfall events may be as, or more, important than changes in rainfall amount in affecting soil CO₂ flux and other components of the carbon cycle highlights the complex nature of ecosystem responses to climate change in North American grasslands.     

内蒙古草甸草原与典型草原地下生物量与生产力季节动态及其碳库潜力

地下根系是草原生态系统的重要组成部分,其生物量及其净生产力对地下碳库具有直接与间接作用,分析地下生物量季节动态与周转对深入揭示草原生态系统碳库动态及其固碳速率与潜力具有重要意义。应用钻土芯法对不同利用方式或管理措施下内蒙古草甸草原、典型草原地下生物量动态及其与温度、降水的相关性研究表明:草甸草原和典型草原地上生物量季节动态均为单峰型曲线,与上月降水显著正相关(P0.05),但地下生物量季节动态表现为草甸草原呈"S"型曲线,典型草原则是双峰型曲线,与温度、降水相关性均不显著(P0.05);两种草原根冠比和地下生物量垂直分布均为指数函数曲线,根茎型草原地下生物量集中在土壤0—5 cm,丛生型草原地下生物量集中于土壤5—10 cm,根冠比值在生长旺季(7—8月份)最小。草甸草原地下净生产力及碳储量范围分别为2167—2953 g m-2a-1和975—1329 gC m-2a-1,典型草原为2342—3333 g m-2a-1和1054—1450 gC m-2a-1,地下净生产力及其碳储量约为地上净生产力及其碳储量的10倍,具有较大的年固碳能力,且相对稳定;地下净生产力与地上净生产力呈显著负相关性(P0.05);地下生物量碳库是地上生物量碳库的10倍左右,适度放牧可增加地下生产力,但长期过度放牧显著降低其地下生物量与生产力,并使其垂直分布趋向于浅层化。     

Growth responses of African savanna trees implicate atmospheric [CO2] as a driver of past and current changes in savanna tree cover

Atmospheric CO₂ has more than doubled since the last glacial maximum (LGM) and could double again within this century, largely due to anthropogenic activity. It has been suggested that low [CO₂] contributed to reduced tree cover in savanna and grassland biomes at LGM, and that increasing [CO₂] over the last century promoted increases in woody plants in these ecosystems over the past few decades. Despite the implications of this idea for understanding global carbon cycle dynamics and key global role of the savanna biome, there are still very few experimental studies quantifying the effects of CO₂ on tree growth and demography in savannas and grasslands. In this paper we present photosynthetic, growth and carbon allocation responses of African savanna trees (Acacia karroo and Acacia nilotica) and a C₄ grass, Themeda triandra, exposed to a gradient of CO₂ concentrations from 180 (typical of LGM) to 1000 µmol mol^?1 in open‐top chambers in a glasshouse as a first empirical test of this idea. Photosynthesis, total stem length, total stem diameter, shoot dry weight and root dry weight of the acacias increased significantly across the CO₂ gradient, saturating at higher CO₂ concentrations. After clipping to simulate fire, plants showed an even greater response in total stem length, total stem diameter and shoot dry weight, signalling the importance of re‐sprouting following disturbances such as fire or herbivory in savanna systems. Root starch (per unit root mass and total root starch per plant) increased steeply along the CO₂ gradient, explaining the re‐sprouting response. In contrast to the strong response of tree seedlings to the CO₂ gradient, grass productivity showed little variation, even at low CO₂ concentrations. These results suggest that CO₂ has significant direct effects on tree recruitment in grassy ecosystems, influencing the ability of trees to recover from fire damage and herbivory. Fire and herbivore regimes that were effective in controlling tree increases in grassy ecosystems could thus be much less effective in a CO₂‐rich world, but field‐based tests are needed to confirm this suggestion.     

青藏高原不同牧草人工草地对土壤线虫群落的影响

为缓解草蓄矛盾,青藏高原人工草地得到快速发展。土壤线虫对环境变化敏感,是草地生态系统的重要组成部分。然而,不同牧草人工草地对土壤线虫群落的影响尚不明确。2016年7月,对多年生禾本科(垂穗披碱草Elymus nutans Griseb.、老芒麦Elymus sibiricus L.、早熟禾Poa annua L.和羊茅Festuca ovina L.)、一年生禾本科燕麦Avena sativa L.和多年生豆科紫花苜蓿Medicago sativa L.等6种单播牧草人工草地(建植期4年)和天然草地(对照)的土壤线虫群落进行了调查。结果表明：(1)土壤线虫隶属于2纲8目32科58属,平均密度为1754个/100 g干土;紫花苜蓿样地的线虫密度最低,为949个/100 g干土;燕麦样地最高,为3267个/100 g干土;(2)与天然草地相比,燕麦样地的线虫群落总密度、多样性以及植食性和杂食-捕食性线虫密度显著增加,而其他人工草地的线虫群落密度、多样性以及植食性、食真菌和食细菌线虫密度均无显著变化;(3)土壤线虫总密度以及各营养类群密度在不同人工草地间差异显著,且均在燕麦样地最高;(4)建...     

Estimating net primary productivity of grasslands from field biomass measurements in temperate northern China

Data on field biomass measurements in temperate grasslands of northern China (141 samples from 74 sites) were obtained from 23 Chinese journals, reports and books. Net primary productivity (NPP) of grasslands was estimated using three algorithms (peak live biomass, peak standing crop and maximum minus minimum live biomass), respectively, based on availability of biomass data in sites. 135 samples which have aboveground biomass (AGB) measurements, have peak AGB ranges from 20 to 2021 g m^–2 (mean = 325.3) and the aboveground NPP (ANPP) ranges from 15 to 1647.1 g m^–2 per year (mean = 295.7). 72 samples which have belowground biomass (BGB) measurements, have peak BGB ranges from 226.5 to 12827.5 g m^–2 (mean = 3116) and the belowground NPP (BNPP) ranges from 15.8 to 12827.5 g m^–2 per year (mean = 2425.6). In total 66 samples have the total NPP (TNPP), ranging from 55.3 to 13347.8 g m^–2 per year (mean = 2980.3). Mean peak biomass and NPP varied from different geographical sampling locations, but they had a general rough regularity in ten grasslands. Meadow, mountain and alpine grasslands had high biomass and NPP (sometimes including saline grassland). Forested steppe, saline grassland and desert had median values. Meadowed and typical steppes had low biomass and NPP (sometimes including desert). The lowest biomass and NPP occurred in deserted steppe and stepped desert. Grassland ANPP has significant positive relationships with annual and summer precipitation as well as summer temperature (all p<0.01). However, grassland BNPP and TNPP have more significant negative relationships with summer temperature (p<0.01) than with annual temperature (p<0.05). The analysis of climate – productivity correlations implied that aboveground productivity is more controlled by rainfall, whereas belowground and total productivity is more influenced by temperature in the temperate grasslands of northern China. The present study might underestimate grassland NPP in northern China due to limitation of biomass measurements. Data on relative long-term aboveground and belowground biomass dynamics, as well as data of standing dead matter, litterfall, decomposition and turnover, are required if grassland NPP is to be more accurately estimated and the role of temperate grasslands in the regional to global carbon cycles is to be fully appreciated. This revised version was published online in June 2006 with corrections to the Cover Date.     

The sensitivity of annual grassland carbon cycling to the quantity and timing of rainfall

Global climate models predict significant changes to the rainfall regimes of the grassland biome, where C cycling is particularly sensitive to the amount and timing of precipitation. We explored the effects of both natural interannual rainfall variability and experimental rainfall additions on net C storage and loss in annual grasslands. Soil respiration and net primary productivity (NPP) were measured in treatment and control plots over four growing seasons (water years, or WYs) that varied in wet‐season length and the quantity of rainfall. In treatment plots, we increased total rainfall by 50% above ambient levels and simulated one early‐ and one late‐season storm. The early‐ and late‐season rain events significantly increased soil respiration for 2–4 weeks after wetting, while augmentation of wet‐season rainfall had no significant effect. Interannual variability in precipitation had large and significant effects on C cycling. We observed a significant positive relationship between annual rainfall and aboveground NPP across the study (P=0.01, r²=0.69). Changes in the seasonal timing of rainfall significantly affected soil respiration. Abundant rainfall late in the wet season in WY 2004, a year with average total rainfall, led to greater net ecosystem C losses due to a ～50% increase in soil respiration relative to other years. Our results suggest that C cycling in annual grasslands will be less sensitive to changes in rainfall quantity and more affected by altered seasonal timing of rainfall, with a longer or later wet season resulting in significant C losses from annual grasslands.     

Plant height as a simple predictor of the root to shoot ratio: Evidence from alpine grasslands on the Tibetan Plateau

Question: Since increases in altitude and grazing intensity generally result in decreases in height growth of alpine grasslands, plant height may integrate effects of environmental stress and grazing disturbance and provide better assessments of the variation in root: shoot (R: S) biomass ratio than other variables. However, it is unclear if there is a general relationship between plant height and R:S ratio across grassland ecosystems. Such knowledge would be helpful for root biomass estimation in grasslands. Location: An altitudinal transect in the Gonghe Basin (2880–4040 m a.s.l.), northeast Tibetan plateau. Methods: We measured standing biomass both above‐ground and below‐ground, maximum plant height (MPH) and soil variables across 43 plots. Results: Climatic variables explained the variations in MPH and R: S ratio of undegraded grasslands better than soil variables (46–50% vs < 19%), while those of degraded grasslands generally showed insignificant correlations with climatic and soil variables. There was a general relationship between R: S ratio and MPH (negative, R²= 0.76, P< 0.001) across degraded and undegraded grasslands. The relationship was used to predict R: S ratio in 13 additional plots in steppe grasslands of Inner Mongolia, and good agreement of expected and observed values has been found (R²= 0.87, P < 0.001). Conclusions: MPH, that is relatively easy to measure, can be used to predict R:S ratio at plot to regional scales. It is promising to develop a new method for large‐scale estimation of root biomass in grasslands using MPH and shoot biomass avoiding tedious procedures of physical measuring of above and below‐ground biomass.     

Regional Variations in Biomass Distribution in Brazilian Savanna Woodland

The Cerrado, the savanna biome in central Brazil, mostly comprised of woodland savanna, is experiencing intense and fast land use changes. To understand the changes in Cerrado carbon stocks, we present an overview of biomass distribution in different Cerrado vegetation types (i.e., grasslands, shrublands and forestlands). We surveyed 26 studies including 170 Cerrado sites. The grasslands presented mean total biomass of 24 Mg/ha, with 70 percent allocated in the belowground portion. In shrublands, the mean total biomass was 58 Mg/ha being 58 percent in the belowground portion. Finally, in forestlands the mean total biomass was 98 Mg/ha with 18 percent as belowground biomass. The surveyed studies presented 12 allometric equations for biomass estimate, most involving both diameter and height. Data on wood density for Cerrado shrubs and trees are not abundant and the average value was 0.66 g/cm³, similar to that found in the central portion of the Amazon Forest. We also examined the relationship between total precipitation and dry‐season intensity with biomass variation in the Cerrado shrubland using data from tropical rainfall measurement mission (TRMM) for the period 2000–2010. Dry‐season precipitation amount in cerrado areas in severe drought regions explained 29 percent of the variation in aboveground woody biomass. This finding is important in the face of the predictions of longer and more severe dry seasons in the region due to climate change.     

Drought effects on above- and belowground production of a grazed temperate grassland ecosystem

The effect of climatic variation on terrestrial aboveground productivity (ANPP) has been well studied. However, little is known about how variable climate, including drought, may influence belowground productivity (BNPP), which constitutes most of the annual primary production of grasslands. The objectives of this study were to (1) examine how a 3-year period of declining moisture, which began as climatically wet to average across Yellowstone National Park (YNP) and ended in drought, affected ANPP and BNPP in grasslands of YNP and (2) how herds of grazing ungulates, which were shown previously to stimulate grassland shoot and root growth in YNP, may have interacted with climatic conditions to influence grassland production. ANPP and 0–20 cm BNPP, representing the bulk of the root dynamics, were measured in grazed and ungrazed (fenced) grassland at nine sites ranging widely in elevation, soil conditions and plant production during the 3-year study. Results revealed that 0–20 cm BNPP was strongly influenced by drought (P = 0.0005) and declined from 1999 to 2001 among ungrazed and grazed grasslands by 39 and 49%, respectively. The greater reduction in 0–20 cm BNPP among grazed grasslands was due, in part, to a decline (P = 0.07) in the stimulatory effect of grazing, i.e., the ratio g BNPP stimulated: g shoot consumed. In contrast, ANPP was unaffected by drought in either type of grassland. Thus, the effect of this drought in YNP was a large reduction in BNPP, which was a function of (1) a direct negative influence of increased moisture stress on root growth and (2) a weak interaction between drought and herbivory that led to a decline in the positive feedback from grazers to BNPP. These findings highlight the need to better understand factors that control root growth and to study the effects of climatic variation on grasslands within an ecosystem framework to include potentially important climate–consumer interactions.