Diverse mechanisms for CO2 effects on grassland litter decomposition

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) can potentially alter litter decomposition rates by changing: (i) the litter quality of individual species, (ii) allocation patterns of individual species, (iii) the species composition of ecosystems (which could alter ecosystem‐level litter quality and allocation), (iv) patterns of soil moisture, and (v) the composition and size of microbial communities. To determine the relative importance of these mechanisms in a California annual grassland, we created four mixtures of litter that differed in species composition (the annual legume Lotus wrangelianus Fischer & C. Meyer comprised either 10% or 40% of the initial mass) and atmospheric [CO₂] during growth (ambient or double‐ambient). These mixtures decomposed for 33 weeks at three positions (above, on, and below the soil surface) in four types of grassland microcosms (fertilized and unfertilized microcosms exposed to elevated or ambient [CO₂]) and at a common field site. Initially, legume‐rich litter mixtures had higher nitrogen concentrations ([N]) than legume‐poor mixtures. In most positions and environments, the different litter mixtures decomposed at approximately the same rate. Fertilization and CO₂ enrichment of microcosms had no effect on mass loss of litter within them. However, mass loss was strongly related to litter position in both microcosms and the field. Nitrogen dynamics of litter were significantly related to the initial [N] of litter on the soil surface, but not in other positions. We conclude that changes in allocation patterns and species composition are likely to be the dominant mechanisms through which ecosystem‐level decomposition rates respond to increasing atmospheric [CO₂].     

Elevated CO2 effects on decomposition processes in a grazed grassland

The effects of elevated atmospheric CO₂ (475 μL L^?1) on in situ decomposition of plant litter and animal faecal material were studied over 2 years in a free air CO₂ enrichment (FACE) facility. The pasture was grazed by sheep and contained a mixture of C₃ and C₄ grasses, legumes and forbs. There was no effect of elevated CO₂ on decomposition within plant species but marked differences between species with faster decomposition in dicots; a group that increased in abundance at elevated CO₂. Decomposition of mixed herbage root material occurred at a similar rate to that of leaf litter suggesting that any CO₂‐induced increase in carbon allocation to roots would not reduce rates of decomposition. Sheep faeces resulting from a ‘high‐CO₂ diet’ decomposed significantly slower during summer but not during winter. The overall outcome of these experiments were explored using scenarios that took account of changes in botanical composition, allocation to roots and the presence of herbivores. In the absence of herbivores, elevated CO₂ led to a 15% increase in the rate of mass loss and an 18% increase in the rate of nitrogen (N) release. In the presence of herbivores, these effects were partially removed (11% increase in rate of mass loss and 9% decrease in N release rate) because of the recycling occurring through the animals in the form of faeces.     

Biomass and compositional changes occur in chalk grassland turves exposed to elevated CO2 for two seasons in FACE

Artificial turves composed of 7 chalk grassland species (Festuca ovina L.; Briza media L.; Bromopsis erecta (Hudson) Fourr.; Plantago media L.; Sanguisorba minor Scop.; Anthyllis vulneraria L. and Lotus corniculatus L.) were grown from seed and exposed to two seasons of elevated (600 μmol mol^–1) and ambient (340 μmol mol^–1) CO₂ concentrations in free air CO₂ enrichment (ETH-FACE, Zurich). The turves were clipped regularly to a height of 5 cm and assessed for above ground biomass production and relative abundance based on accumulated clipped dry biomass as well as by point quadrat recording. Below ground biomass production was assessed with root in-growth bags during the second season of growth. Increases in total biomass (> 30%) were noted in elevated CO₂, but the differences did not become significant until the second season of growth. Individual species’ biomass varied in response to elevated CO₂, with significant increases in biomass in elevated CO₂ turves for both legume species, and no significant CO₂ effect on S. minor or P. media. An initial positive CO₂ effect on biomass of combined grass species was reversed by the end of the experiment with less biomass and a significantly smaller proportion of total biomass present in elevated CO₂, which was attributed primarily to changes in proportion of F. ovina. Species relative abundance was significantly affected by elevated CO₂ in the final 4 of the 6 clip events, with the legume species increasing in proportion at the expense of the other species, particularly the grasses. Root length and dry weight were both significantly increased in elevated CO₂ (77% and 89%, respectively), and these increases were greater than increases in shoot biomass (36%) from the same period. Species responses to elevated CO₂, within the model community, were not consistent with predictions made from data on individual species, leading to the conclusion that responses to elevated CO₂, at the community level, and species within the community level, are the result of direct physiological effects and indirect competitive effects. These conclusions are discussed with respect to the ecological responses of natural communities, and the chalk grassland community in particular, to elevated CO₂.     

Legumes regulate grassland soil N cycling and its response to variation in species diversity and N supply but not CO2

Legumes are an important component of plant diversity that modulate nitrogen (N) cycling in many terrestrial ecosystems. Limited knowledge of legume effects on soil N cycling and its response to global change factors and plant diversity hinders a general understanding of whether and how legumes broadly regulate the response of soil N availability to those factors. In a 17‐year study of perennial grassland species grown under ambient and elevated (+180 ppm) CO₂ and ambient and enriched (+4 g N m^?2 year^?1) N environments, we compared pure legume plots with plots dominated by or including other herbaceous functional groups (and containing one or four species) to assess the effect of legumes on N cycling (net N mineralization rate and inorganic N pools). We also examined the effects of numbers of legume species (from zero to four) in four‐species mixed plots on soil N cycling. We hypothesized that legumes would increase N mineralization rates most in those treatments with the greatest diversity and the greatest relative limitation by and competition for N. Results partially supported these hypotheses. Plots with greater dominance by legumes had greater soil nitrate concentrations and mineralization rates. Higher species richness significantly increased the impact of legumes on soil N metrics, with 349% and 505% higher mineralization rates and nitrate concentrations in four‐species plots containing legumes compared to legume‐free four‐species plots, in contrast to 185% and 129% greater values, respectively, in pure legume than nonlegume monoculture plots. N‐fertilized plots had greater legume effects on soil nitrate, but lower legume effects on net N mineralization. In contrast, neither elevated CO₂ nor its interaction with legumes affected net N mineralization. These results indicate that legumes markedly influence the response of soil N cycling to some, but not all, global change drivers.     

Plant species richness, elevated CO2, and atmospheric nitrogen deposition alter soil microbial community composition and function

We determined soil microbial community composition and function in a field experiment in which plant communities of increasing species richness were exposed to factorial elevated CO₂ and nitrogen (N) deposition treatments. Because elevated CO₂ and N deposition increased plant productivity to a greater extent in more diverse plant assemblages, it is plausible that heterotrophic microbial communities would experience greater substrate availability, potentially increasing microbial activity, and accelerating soil carbon (C) and N cycling. We, therefore, hypothesized that the response of microbial communities to elevated CO₂ and N deposition is contingent on the species richness of plant communities. Microbial community composition was determined by phospholipid fatty acid analysis, and function was measured using the activity of key extracellular enzymes involved in litter decomposition. Higher plant species richness, as a main effect, fostered greater microbial biomass, cellulolytic and chitinolytic capacity, as well as the abundance of saprophytic and arbuscular mycorrhizal (AM) fungi. Moreover, the effect of plant species richness on microbial communities was significantly modified by elevated CO₂ and N deposition. For instance, microbial biomass and fungal abundance increased with greater species richness, but only under combinations of elevated CO₂ and ambient N, or ambient CO₂ and N deposition. Cellobiohydrolase activity increased with higher plant species richness, and this trend was amplified by elevated CO₂. In most cases, the effect of plant species richness remained significant even after accounting for the influence of plant biomass. Taken together, our results demonstrate that plant species richness can directly regulate microbial activity and community composition, and that plant species richness is a significant determinant of microbial response to elevated CO₂ and N deposition. The strong positive effect of plant species richness on cellulolytic capacity and microbial biomass indicate that the rates of soil C cycling may decline with decreasing plant species richness.     

Leaf gas exchange and nitrogen dynamics of N2-fixing, field-grown Alnus glutinosa under elevated atmospheric CO2

Few studies have investigated the effects of elevated CO₂ on the physiology of symbiotic N₂-fixing trees. Tree species grown in low N soils at elevated CO₂ generally show a decline in photosynthetic capacity over time relative to ambient CO₂ controls. This negative adjustment may be due to a reallocation of leaf N away from the photosynthetic apparatus, allowing for more efficient use of limiting N. We investigated the effect of twice ambient CO₂ on net CO₂ assimilation (A), photosynthetic capacity, leaf dark respiration, and leaf N content of N2-fixing Alnus glutinosa (black alder) grown in field open top chambers in a low N soil for 160 d. At growth CO₂, A was always greater in elevated compared to ambient CO₂ plants. Late season A vs. internal leaf p(CO₂) response curves indicated no negative adjustment of photosynthesis in elevated CO₂ plants. Rather, elevated CO₂ plants had 16% greater maximum rate of CO₂ fixation by Rubisco. Leaf dark respiration was greater at elevated CO₂ on an area basis, but unaffected by CO₂ on a mass or N basis. In elevated CO₂ plants, leaf N content (μg N cm^?2) increased 50% between Julian Date 208 and 264. Leaf N content showed little seasonal change in ambient CO₂ plants. A single point acetylene reduction assay of detached, nodulated root segments indicated a 46% increase in specific nitrogenase activity in elevated compared to ambient CO₂ plants. Our results suggest that N₂-fixing trees will be able to maintain high A with minimal negative adjustment of photosynthetic capacity following prolonged exposure to elevated CO₂ on N-poor soils.     

Selective grazing modifies previously anticipated responses of plant community composition to elevated CO2 in a temperate grassland

Our limited understanding of terrestrial ecosystem responses to elevated CO₂ is a major constraint on predicting the impacts of climate change. A change in botanical composition has been identified as a key factor in the CO₂ response with profound implications for ecosystem services such as plant production and soil carbon storage. In temperate grasslands, there is a strong consensus that elevated CO₂ will result in a greater physiological stimulus to growth in legumes and to a lesser extent forbs, compared with C3 grasses, and the presumption this will lead in turn to a greater proportion of these functional groups in the plant community. However, this view is based on data mainly collected in experiments of three or less years in duration and not in experiments where defoliation has been by grazing animals. Grazing is, however, the most common management of grasslands and known in itself to influence botanical composition. In a long‐term Free Air Carbon Dioxide Enrichment (FACE) experiment in a temperate grassland managed with grazing animals (sheep), we found the response to elevated CO₂ in plant community composition in the first 5 years was consistent with the expectation of increased proportions of legumes and forbs. However, in the longer term, these differences diminished so that the proportions of grasses, legumes and forbs were the same under both ambient and elevated CO₂. Analysis of vegetation before and after each grazing event showed there was a sustained disproportionately greater removal (‘apparent selection’) of legumes and forbs by the grazing animals. This bias in removal was greater under elevated CO₂ than ambient CO₂. This is consistent with sustained faster growth rates of legumes and forbs under elevated CO₂ being countered by selective defoliation, and so leading to little difference in community composition.     

Effect of CO2 enrichment on non-structural carbohydrates in leaves, stems and ears of spring wheat

Field-grown spring wheat (Triticum aestivum L. cv. Dragon) was exposed to ambient and elevated CO₂ concentrations (1.5 and 2 times ambient) in open-top chambers. Contents of non-structural carbohydrates were analysed enzymatically in leaves, stems and ears six times during the growing season. The impact of elevated CO₂ on wheat carbohydrates was non-significant in most harvests. However, differences in the carbohydrate contents due to elevated CO₂ were found in all plant compartments. Before anthesis, at growth stage (GS) 30 (the stem is 1 cm to the shoot apex), the plants grown in elevated CO₂ contained significantly more water soluble carbohydrates (WSC), fructans, starch and total non-structural carbohydrates (TNC) in the leaves in comparison with the plants grown in ambient CO₂. It is hypothesised that the plants from the treatments with elevated CO₂ were sink-limited at GS30. After anthesis, the leaf WSC and TNC contents of the plants from elevated CO₂ started to decline earlier than those of the plants from ambient CO₂. This may indicate that the leaves of plants grown in the chambers with elevated CO₂ senesced earlier. Elevated CO₂ accelerated grain development: 2 weeks after anthesis, the plants grown in elevated CO₂ contained significantly more starch and significantly less fructans in the ears compared to the plants grown in ambient CO₂. Elevated CO₂ had no effect on ear starch and TNC contents at the final harvest. Increasing the CO₂ concentration from 360 to 520 μmol mol^?1 had a larger effect on wheat non-structural carbohydrates than the further increase from 520 to 680 μmol mol^?1. The results are discussed in relation to the effects of elevated CO₂ on yield and yield components.     

Mineral nitrogen cycling through earthworm casts in a grazed pasture under elevated atmospheric CO2

We used the New Zealand grazed pasture free air CO₂ enrichment experiment to determine the effects of elevated CO₂ on earthworm (Aporrectodea caliginosa and Lumbricus rubellus) cast production and mineral nitrogen (N) concentration over a 5‐week period in the spring. Elevated CO₂ did not affect earthworm biomass or the amount of cast material produced. However, cast mineral N concentrations were 18% lower, resulting in 27% less mineral N being deposited on the soil surface under elevated CO₂. An analysis of the earthworms' potential diet showed that a reduction in the N content of sheep dung at elevated CO₂ was the most likely cause of the lower cast N concentrations. Earthworm casts made only a small contribution to mineral N cycling in our system; however, their quality may act as a sensitive indicator of reduced N availability under elevated CO₂ which is consistent with the hypothesised process of progressive N limitation.     

Altered physiological and growth responses to elevated [CO2] in offspring from holm oak (Quercus ilex L.) mother trees with lifetime exposure to naturally elevated [CO2]

An important question with respect to plant performance in future climatic scenarios is whether the offspring of mature trees that have experienced lifelong exposure to elevated [CO₂] show altered physiological responses to elevated [CO₂] compared with those originating from current ambient CO₂ concentrations. To investigate this question, acorns were collected from two seed sources, denoted as ‘control’ and ‘spring’, from Quercus ilex mother trees grown at ambient (36 Pa) and at about twice ambient CO₂ concentrations, respectively, close to a natural CO₂ spring, Laiatico, central Italy. The seedlings were raised for 8 months under controlled conditions at ambient and elevated [CO₂] in a reciprocal experimental design and were used for the determination of biomass, photosynthesis and foliar carbohydrate concentrations, as well as the accumulation of structural biomass and lignin during leaf maturation. Under ambient [CO₂], biomass and foliar carbon acquisition in control progeny were not significantly different from spring progeny. However, under elevated [CO₂], spring seedlings showed less CO₂ acclimation than control seedlings but no significant differences in non‐structural carbohydrate concentrations and structural biomass per unit leaf dry mass. Developmental lignin accumulation in leaves was delayed under elevated [CO₂] compared with ambient [CO₂], but only in control progeny. Under elevated [CO₂], whole‐plant biomass, leaf area and stem diameter were significantly increased in Quercus ilex seedlings from both seed sources but with a higher stimulation of above‐ground biomass in spring than in control seedlings and a higher stimulation of below‐ground biomass in control seedlings. These results indicate that life history and/or progeny may determine the species‐specific CO₂ response and suggest that positive CO₂ acclimation is possible.