Effects of age and ontogeny on photosynthetic responses of a determinate annual plant to elevated CO2 concentrations

Plant responses to elevated CO₂ concentrations ([CO₂]) may be regulated by both accelerated ontogeny and allocational changes as plants grow. However, isolating ontogeny‐related effects from age‐related effects are difficult because these factors are often confounded. In this study, the roles of age and ontogeny in photosynthetic responses to elevated [CO₂] were examined on Xanthium strumarium L. grown at ambient (365 µmol mol^?1) and elevated (730 µmol mol^?1) [CO₂]. To examine age‐related effects, six cohorts were planted at 5‐day intervals. To examine ontogeny‐related effects, all plants were induced to flower at the same time; ontogeny in Xanthium is relatively unaffected by growth in elevated [CO₂]. Growth in elevated [CO₂] increased net photosynthetic rates by approximately 30% throughout vegetative growth (i.e. active carbohydrate sinks), approximately 10% during flowering (i.e. minimal sink activity), and approximately 20% during fruit production (i.e. active sinks). At the harvest, the ratio of source to sink tissue significantly decreased with increasing plant age and was correlated with leaf soluble sugar concentration. Leaf soluble sugar concentration was negatively correlated with the relative photosynthetic response to elevated [CO₂]. These results suggest that age and ontogeny independently affect photosynthetic responses to elevated [CO₂] and the effects are mediated by reversible changes in source : sink balance.     

Response of plant roots to elevated atmospheric carbon dioxide

Plant root response to atmospheric CO₂ enrichment can be great. Results from this controlled environment investigation demonstrate substantial effects on root system architecture, micromorphology and physiology. The most pronounced effects were an increase in root length (110%) and root dry weight (143%). Root diameter, stele diameter, cortex width, root/shoot and root weight ratios all increased; root numbers did not increase. The long-term implications for belowground processes could be enormous.     

Rice production in a changing climate: a meta-analysis of responses to elevated carbon dioxide and elevated ozone concentration

Rice is arguably the most important food source on the planet and is consumed by over half of the world's population. Considerable increases in yield are required over this century to continue feeding the world's growing population. This meta-analysis synthesizes the research to date on rice responses to two elements of global change, rising atmospheric carbon dioxide concentration ([CO₂]) and rising tropospheric ozone concentration ([O₃]). On an average, elevated [CO₂] (627 ppm) increased rice yields by 23%. Modest increases in grain mass and larger increases in panicle and grain number contributed to this response. The response of rice to elevated [CO₂] varied with fumigation technique. The more closely the fumigation conditions mimicked field conditions, the smaller was the stimulation of yield by elevated [CO₂]. Free air concentration enrichment (FACE) experiments showed only a 12% increase in rice yield. The rise in atmospheric [CO₂] will be accompanied by increases in tropospheric O₃ and temperature. When compared with rice grown in charcoal-filtered air, rice exposed to 62 ppb O₃ showed a 14% decrease in yield. Many determinants of yield, including photosynthesis, biomass, leaf area index, grain number and grain mass, were reduced by elevated [O₃]. While there have been too few studies of the interaction of CO₂ and O₃ for meta-analysis, the interaction of temperature and CO₂ has been studied more widely. Elevated temperature treatments negated any enhancement in rice yield at elevated [CO₂], which suggests that identifying high temperature tolerant germplasm will be key to realizing yield benefits in the future.     

Issues and perspectives for investigating root responses to elevated atmospheric carbon dioxide

A thorough assessment of how plants and ecosystems will respond to increasing concentrations of atmospheric CO₂ requires that the responses of root systems and associated belowground processes be understood. Static measures of root-to-shoot ratio have not been satisfactory for describing the integrated responses of plants to CO₂-enriched atmospheres, but research with a process orientation has suggested that elevated CO₂ can stimulate root growth or root activity and provide a positive feedback on plant growth. There are, however, critical questions concerning the relevance of root data from short-term studies with potted plants when scaling to questions about plants in the field. Data on root responses to CO₂ enrichment in the field are fragmentary, but they allow us to more clearly define research questions for further investigation. Three perspectives for analyzing the significance of root responses as a component of the overall response of the terrestrial biosphere to increasing atmospheric CO₂ are suggested: (1) roots as a platform for nutrient acquisition and a mediator of whole-plant response to CO₂; (2) carbon storage in roots as a component of whole-plant carbon storage; and (3) effects of CO₂ enrichment on root turnover and the implications for carbon storage as soil organic matter. The relative importance of these different perspectives will vary depending on the ecosystem of interest and the larger-scale issues being considered.     

Root production and demography in a california annual grassland under elevated atmospheric carbon dioxide

This study examined root production and turnover in a California grassland during the third year of a long‐term experiment with ambient (LO) and twice‐ambient atmospheric CO₂ (HI), using harvests, ingrowth cores, and minirhizotrons. Based on one‐time harvest data, root biomass was 32% greater in the HI treatment, comparable to the stimulation of aboveground production during the study year. However, the 30–70% increase in photosynthesis under elevated CO₂ for the dominant species in our system is considerably larger than the combined increase in above and belowground biomass. One possible explanation is, increased root turnover, which could be a sink for the additional fixed carbon. Cumulative root production in ingrowth cores from both treatments harvested at four dates was 2–3 times that in the single harvested cores, suggesting substantial root turnover within the growing season. Minirhizotron data confirmed this result, demonstrating that production and mortality occurred simultaneously through much of the season. As a result, cumulative root production was 54%, 47% and 44% greater than peak standing root length for the no chamber (X), LO, and HI plots, respectively. Elevated CO₂, however, had little effect on rates of turnover (i.e. rates of turnover were equal in the LO and HI plots throughout most of the year) and cumulative root production was unaffected by treatment. Elevated CO₂ increased monthly production of new root length (59%) only at the end of the season (April–June) when root growth had largely ceased in the LO plots but continued in the HI plots. This end‐of‐season increase in production coincided with an 18% greater soil moisture content in the HI plots previously described. Total standing root length was not affected by CO₂ treatment. Root mortality was unaffected by elevated CO₂ in all months except April, in which plants grown in the HI plots had higher mortality rates. Together, these results demonstrate that root turnover is considerable in the grassland community and easily missed by destructive soil coring. However, increased fine root turnover under elevated CO₂ is apparently not a major sink for extra photosynthate in this system.     

New insights into the cellular mechanisms of plant growth at elevated atmospheric carbon dioxide concentrations

Rising atmospheric carbon dioxide concentration ([CO₂]) significantly influences plant growth, development, and biomass. Increased photosynthesis rate, together with lower stomatal conductance, has been identified as the key factors that stimulate plant growth at elevated [CO₂] (e[CO₂]). However, variations in photosynthesis and stomatal conductance alone cannot fully explain the dynamic changes in plant growth. Stimulation of photosynthesis at e[CO₂] is always associated with post‐photosynthetic secondary metabolic processes that include carbon and nitrogen metabolism, cell cycle functions, and hormonal regulation. Most studies have focused on photosynthesis and stomatal conductance in response to e[CO₂], despite the emerging evidence of e[CO₂]'s role in moderating secondary metabolism in plants. In this review, we briefly discuss the effects of e[CO₂] on photosynthesis and stomatal conductance and then focus on the changes in other cellular mechanisms and growth processes at e[CO₂] in relation to plant growth and development. Finally, knowledge gaps in understanding plant growth responses to e[CO₂] have been identified with the aim of improving crop productivity under a CO₂ rich atmosphere.     

Age changes in the vegetative vs. reproductive allocation by module demographic strategies in a perennial plant

Retama sphaerocarpa (L.) Boiss. is aMediterranean shrub with a remarkably simplified metameric structure. Terminalyoungest shoots act as units of modular growth, being able to produce newshootsby basal axillary buds (at the base of the shoot) and inflorescencesby lateral axillary buds. In this study, we have analysed the structural andgrowth potential features of these modules, as well as theirdemographic proportions, regarding the allocation of newvegetative and reproductive growth in plants of different age. Reproductiveeffort is proportionally higher in older plants. This shift in the allocationstrategy with plant ontogeny is not attained with changes in the shoot modules(which maintain a constant size, nutrient composition and show a similarnew growth investment per module) but through a differentdemographic composition of the population ofmodulesaccording to their developmental fate (vegetative or reproductive).This indicates a high level of iterativity and a purely modular growth, sincethe attributes of the individual (age of the plants) do not seem toaffect those of the integrating modular units (growth performance of theshoots).     

Super-optimal temperatures are detrimental to peanut (Arachis hypogaea L.) reproductive processes and yield at both ambient and elevated carbon dioxide

Continuing increases in atmospheric carbon dioxide concentration (CO₂) will likely be accompanied by global warming. Our research objectives were (a) to determine the effects of season‐long exposure to daytime maximum/nighttime minimum temperatures of 32/22, 36/26, 40/30 and 44/34°C at ambient (350 μmol mol^?1) and elevated (700 μmol mol^?1) CO₂ on reproductive processes and yield of peanut, and (b) to evaluate whether the higher photosynthetic rates and vegetative growth at elevated CO₂ will negate the detrimental effects of high temperature on reproductive processes and yield. Doubling of CO₂ increased leaf photosynthesis and seed yield by 27% and 30%, respectively, averaged across all temperatures. There were no effects of elevated CO₂ on pollen viability, seed‐set, seed number per pod, seed size, harvest index or shelling percentage. At ambient CO₂, seed yield decreased progressively by 14%, 59% and 90% as temperature increased from 32/22 to 36/26, 40/30 and 44/34°C, respectively. Similar percentage decreases in seed yield occurred at temperatures above 32/22°C at elevated CO₂ despite greater photosynthesis and vegetative growth. Decreased seed yields at high temperature were a result of lower seed‐set due to poor pollen viability, and smaller seed size due to decreased seed growth rates and decreased shelling percentages. Seed harvest index decreased from 0.41 to 0.05 as temperature increased from 32/22 to 44/34°C under both ambient and elevated CO₂. We conclude that there are no beneficial interactions between elevated CO₂ and temperature, and that seed yield of peanut will decrease under future warmer climates, particularly in regions where present temperatures are near or above optimum.     

Responses of five naturalised ornamental freshwater plant species to elevated carbon dioxide concentration and nutrient enrichment

Hydrobiologia - Ongoing global environmental changes such as rising atmospheric CO2 concentration ([CO2]) and eutrophication are in some parts thought to facilitate exotic plant species invasions...     

Seedling density modifies the growth responses of yellow birch maternal families to elevated carbon dioxide

We studied seedling growth responses to ambient and elevated CO₂ (350 and 700 μL L^?1) of three maternal families of yellow birch (Betula alleghaniensis), raised both individually and in high-density stands. Seedlings in competitive, dense stands exhibited markedly lower average CO₂-induced growth enhancements than individually grown plants (16% vs. 49%). Maternal families differed in their growth responses to elevated CO₂. However, differences among families were contingent upon density; families which exhibited the greatest CO₂-induced growth at low density exhibited the least CO₂-responsiveness at high density. These data are discussed in two separate contexts; the reliability of estimates of the CO² fertilization potential of forest species based solely on individually grown plants, and the potential evolutionary consequences of rising CO₂ on regenerating forest tree populations.     

Common bacterial responses in six ecosystems exposed to 10 years of elevated atmospheric carbon dioxide

Six terrestrial ecosystems in the USA were exposed to elevated atmospheric CO(2) in single or multifactorial experiments for more than a decade to assess potential impacts. We retrospectively assessed soil bacterial community responses in all six-field experiments and found ecosystem-specific and common patterns of soil bacterial community response to elevated CO(2) . Soil bacterial composition differed greatly across the six ecosystems. No common effect of elevated atmospheric CO(2) on bacterial biomass, richness and community composition across all of the ecosystems was identified, although significant responses were detected in individual ecosystems. The most striking common trend across the sites was a decrease of up to 3.5-fold in the relative abundance of Acidobacteria Group 1 bacteria in soils exposed to elevated CO(2) or other climate factors. The Acidobacteria Group 1 response observed in exploratory 16S rRNA gene clone library surveys was validated in one ecosystem by 100-fold deeper sequencing and semi-quantitative PCR assays. Collectively, the 16S rRNA gene sequencing approach revealed influences of elevated CO(2) on multiple ecosystems. Although few common trends across the ecosystems were detected in the small surveys, the trends may be harbingers of more substantive changes in less abundant, more sensitive taxa that can only be detected by deeper surveys. Representative bacterial 16S rRNA gene clone sequences were deposited in GenBank with Accession No. JQ366086–JQ387568.     

Responses of trees to elevated carbon dioxide and climate change

The enhancement in photosynthesis at elevated concentration of carbon dioxide level than the ambient level existing in the atmosphere is widely known. However, many of the earlier studies were based on instantaneous responses of plants grown in pots. The availability of field chambers for growing trees, and long-term exposure studies of tree species to elevated carbon dioxide, has changed much of our views on carbon dioxide acting as a fertiliser. Several tree species showed acclimation or even down-regulation of photosynthetic responses while a few of them showed higher photosynthesis and better growth responses. Whether elevated levels of carbon dioxide can serve as a fertilizer in a changed climate scenario still remains an unresolved question. Forest-Air-Carbon dioxide-Enrichment (FACE) sites monitored at several locations have shown lately, that the acclimation or down regulation as reported in chamber studies is not as wide-spread as originally thought. FACE studies predict that there could be an increase of 23–28% productivity of trees at least till 2050. However, the increase in global temperature could also lead to increased respiration, and limitation of minerals in the soil could lead to reduced responses in growth. Elevated carbon dioxide induces partial closure of leaf stomata, which could lead to reduced transpiration and more economical use of water by the trees. Even if the carbon dioxide acts as a fertilizer, the responses are more pronounced only in young trees. And if there are variations in species responses to growth due to elevated carbon dioxide, only some species are going to dominate the natural vegetation. This will have serious implications on the biodiversity and the structure of the ecosystems. This paper reviews the research done on trees using elevated CO₂ and tries to draw conclusions based on different methods used for the study. It also discusses the possible functional variations in some tree species due to climate change.     

Influence of plant diversity and elevated atmospheric carbon dioxide levels on belowground bacterial diversity

Background  

Changes in aboveground plant species diversity as well as variations of environmental conditions such as exposure of ecosystems to elevated concentrations of atmospheric carbon dioxide may lead to changes in metabolic activity, composition and diversity of belowground microbial communities, both bacterial and fungal.     

Responses of soil biota to elevated atmospheric carbon dioxide

Increasing concentrations of atmospheric CO₂ could have dramatic effects upon terrestrial ecosystems including changes in ecosystem structure, nutrient cycling rates, net primary production, C source-sink relationships and successional patterns. All of these potential changes will be constrained to some degree by below ground processes and mediated by responses of soil biota to indirect effects of CO₂ enrichment. A review of our current state of knowledge regarding responses of soil biota is presented, covering responses of mycorrhizae, N-fixing bacteria and actinomycetes, soil microbiota, plant pathogens, and soil fauna. Emphasis will be placed on consequences to biota of increasing C input through the rhizosphere and resulting feedbacks to above ground systems. Rising CO₂ may also result in altered nutrient concentrations of plant litter, potentially changing decomposition rates through indirect effects upon decomposer communities. Thus, this review will also cover current information on decomposition of litter produced at elevated CO₂. Summary Predictably, the responses of soil biota to CO₂ enrichment and the degree of experimental emphasis on them increase with proximity to, and intimacy with, roots. Symbiotic associations are all stimulated to some degree. Total plant mycorrhization increases with elevated CO₂. VAM fungi increase proportionately with fine root length/mass increase. ECM fungi, however, exhibit greater colonization per unit root length/mass at elevated CO₂ than at current atmospheric levels. Total N-fixation per plant increases in all species examined, although the mechanisms of increase, as well as the eventual benefit to the host relative to N uptake may vary. Microbial responses are unclear. The assumption that changes in root exudation will drive increased mineralization and facilitate nutrient uptake should be examined experimentally, in light of recent models. Microbial results to date suggest that metabolic activity (measured as changes in process rates) is stimulated by root C input, rather than population size (measured by cell or colony counts). Insufficient evidence exists to predict responses of either soil-borne plant pathogens or soil fauna (i.e., food web responses). These are areas requiring attention, the first for its potential to limit ecosystem production through disease and the second because of its importance to nutrient cycling processes. Preliminary data on foliar litter decomposition suggests that neither nutrient ratios nor decomposition rates will be affected by rising CO₂. This is another important area that may be better understood as the number of longer term studies with more realistic CO₂ exposures increase. Evidence continues to mount that C fixation increases with CO₂ enrichment and that the bulk of this C enters the belowground component of ecosystems. The global fate and effects of this additional C may affect all hierarchical levels, from organisms to ecosystems, and will be largely determined by responses of soil biota.     

Plant and fungal responses to elevated atmospheric carbon dioxide in mycorrhizal seedlings of Pinus sylvestris

The effects of elevated CO₂ concentration upon the mycorrhizal relationships of Scots pine (Pinus sylvestris) seedlings were investigated. Plants were grown for 4 months with their shoots exposed to ambient (C_AMB=360 μl l⁻¹) or elevated (C_ELEV=700 μl l⁻¹) CO₂ environments while their root systems, either colonised by the mycorrhizal fungi Paxillus involutus or Suillus bovinus, or left in the non-mycorrhizal condition, were maintained in sealed dishes. In one series of these plants the effects of C_ELEV upon the extent of mycorrhizal development and upon their growth and nutrition were determined, while another series were transferred from the dishes after 1 month, to transparent observation chambers before being returned to the two CO₂ environments. In these chambers, the effects of C_ELEV upon development of the external mycelial systems of the two mycorrhizal fungi was determined by measuring the advance of the hyphal fronts of the mycorrhizal fungi across non-sterile peat from the colonised plants. The dry mass and number of mycorrhizal tips were significantly higher in C_ELEV than in the C_AMB condition in plants colonised by both fungi in the dishes. Yields of whole plants and of shoots were higher in the C_ELEV treatment whether or not they were grown in the mycorrhizal condition, but the greater yields were not associated in these sealed systems with enhanced nutrient gain. The dry mass of non-mycorrhizal plants was greater than that of those colonised by mycorrhizal fungi under elevated CO₂. This is thought to be attributable to the energetic cost of production of the larger mycorrhizal systems in this treatment. The extent of development of the mycorrhizal mycelial systems of both fungi was greatly increased in C_ELEV relative to that in C_AMB environments. It is hypothesised that increased allocation of carbon to mycorrhizal root systems and their associated mycelia would provide the potential for enhancement of nutrient acquisition in open systems of greater fertility.