Role of polyhydroxybutyrate and glycogen as carbon storage compounds in pea and bean bacteroids

Rhizobium leguminosarum synthesizes polyhydroxybutyrate and glycogen as its main carbon storage compounds. To examine the role of these compounds in bacteroid development and in symbiotic efficiency, single and double mutants of R. legumosarum bv. viciae were made which lack polyhydroxybutyrate synthase (phaC), glycogen synthase (glgA), or both. For comparison, a single phaC mutant also was isolated in a bean-nodulating strain of R. leguminosarum bv. phaseoli. In one large glasshouse trial, the growth of pea plants inoculated with the R. leguminosarum bv. viciae phaC mutant were significantly reduced compared with wild-type-inoculated plants. However, in subsequent glasshouse and growth-room studies, the growth of pea plants inoculated with the mutant were similar to wildtype-inoculated plants. Bean plants were unaffected by the loss of polyhydroxybutyrate biosynthesis in bacteroids. Pea plants nodulated by a glycogen synthase mutants or the glgA/phaC double mutant, grew as well as the wild type in growth-room experiments. Light and electron micrographs revealed that pea nodules infected with the glgA mutant accumulated large amounts of starch in the II/III interzone. This suggests that glycogen may be the dominant carbon storage compound in pea bacteroids. Polyhydroxybutyrate was present in bacteria in the infection thread of pea plants but was broken down during bacteroid formation. In nodules infected with a phaC mutant of R. leguminosarum bv. viciae, there was a drop in the amount of starch in the II/III interzone, where bacteroids form. Therefore, we propose a carbon burst hypothesis for bacteroid formation, where polyhydroxybutyrate accumulated by bacteria is degraded to fuel bacteroid differentiation.

Role of polyhydroxybutyrate and glycogen as carbon storage compounds in pea and bean bacteroids

Rhizobium leguminosarum synthesizes polyhydroxybutyrate and glycogen as its main carbon storage compounds. To examine the role of these compounds in bacteroid development and in symbiotic efficiency, single and double mutants of R. legumosarum bv. viciae were made which lack polyhydroxybutyrate synthase (phaC), glycogen synthase (glgA), or both. For comparison, a single phaC mutant also was isolated in a bean-nodulating strain of R. leguminosarum bv. phaseoli. In one large glasshouse trial, the growth of pea plants inoculated with the R. leguminosarum bv. viciae phaC mutant were significantly reduced compared with wild-type-inoculated plants. However, in subsequent glasshouse and growth-room studies, the growth of pea plants inoculated with the mutant were similar to wildtype-inoculated plants. Bean plants were unaffected by the loss of polyhydroxybutyrate biosynthesis in bacteroids. Pea plants nodulated by a glycogen synthase mutants or the glgA/phaC double mutant, grew as well as the wild type in growth-room experiments. Light and electron micrographs revealed that pea nodules infected with the glgA mutant accumulated large amounts of starch in the II/III interzone. This suggests that glycogen may be the dominant carbon storage compound in pea bacteroids. Polyhydroxybutyrate was present in bacteria in the infection thread of pea plants but was broken down during bacteroid formation. In nodules infected with a phaC mutant of R. leguminosarum bv. viciae, there was a drop in the amount of starch in the II/III interzone, where bacteroids form. Therefore, we propose a carbon burst hypothesis for bacteroid formation, where polyhydroxybutyrate accumulated by bacteria is degraded to fuel bacteroid differentiation.

A case study to investigate the life cycle carbon emissions and carbon storage capacity of a cross laminated timber, multi-storey residential building

Forests are a store of carbon and an eco-system that continually removes carbon dioxide from the atmosphere. If they are sustainably managed, the carbon store can be maintained at a constant level, while the trees removed and converted to timber products can form an additional long term carbon store. The total carbon store in the forest and associated ‘wood chain’ therefore increases over time, given appropriate management. This increasing carbon store can be further enhanced with afforestation. The UK’s forest area has increased continually since the early 1900s, although the rate of increase has declined since its peak in the late 1980s, and it is a similar picture in the rest of Europe. The increased sustainable use of timber in construction is a key market incentive for afforestation, which can make a significant contribution to reducing carbon emissions. The case study presented in this paper demonstrates the carbon benefits of a Cross Laminated Timber (CLT) solution for a multi-storey residential building in comparison with a more conventional reinforced concrete solution. The embodied carbon of the building up to completion of construction is considered, together with the stored carbon during the life of the building and the impact of different end of life scenarios. The results of the study show that the total stored carbon in the CLT structural frame is 1215tCO2 (30tCO2 per housing unit). The choice of treatment at end of life has a significant effect on the whole life embodied carbon of the CLT frame, which ranges from -1017 tCO2e for re-use to +153tCO2e for incinerate without energy recovery. All end of life scenarios considered result in lower total CO2e emissions for the CLT frame building compared with the reinforced concrete frame solution.

Reforesting degraded agricultural landscapes with Eucalypts: effects on carbon storage and soil fertility after 26years

In the Western Australian wheatbelt, the restoration of native eucalypt forests for managing degraded agricultural landscapes is a critical part of managing dryland salinity and rebuilding biodiversity. Such reforestation will also sequester carbon. Whereas most investigative emphasis has been on carbon stored in biomass, the effects of reforestation on soil organic carbon (SOC) stores and fertility are not known. Two 26 year old reforestation experiments with four Eucalyptus species (E. cladocalyx var nana, E. occidentalis, E. sargentii and E. wandoo) were compared with agricultural sites (Field). SOC stores (to 0.3 m depth) ranged between 33 and 55 Mg ha−1, with no statistically significant differences between tree species and adjacent farmland. Farming comprised crop and pasture rotations. In contrast, the reforested plots contained additional carbon in the tree biomass (23–60 Mg ha−1) and litter (19–34 Mg ha−1), with the greatest litter accumulation associated with E. sargentii. Litter represented between 29 and 56% of the biomass carbon and the protection or utilization of this litter in fire-prone, semi-arid farmland will be an important component of carbon management. Exch-Na and Exch-Mg accumulated under E. sargentii and E. occidentalis at one site. The results raise questions about the conclusions of SOC sequestration studies following reforestation based on limited sampling and reiterate the importance of considering litter in reforestation carbon accounts.

Cropping systems, carbon sequestration and erosion in Brazil, a review

Soils represent a large carbon pool, approximately 1500 Gt, which is equivalent to almost three times the quantity stored in terrestrial biomass and twice the amount stored in the atmosphere. Any modification of land use or land management can induce variations in soil carbon stocks, even in agricultural systems that are perceived to be in a steady state. Tillage practices often induce soil aerobic conditions that are favourable to microbial activity and may lead to a degradation of soil structure. As a result, mineralisation of soil organic matter increases in the long term. The adoption of no-tillage systems and the maintenance of a permanent vegetation cover using Direct seeding Mulch-based Cropping system or DMC, may increase carbon levels in the topsoil. In Brazil, no-tillage practices (mainly DMC), were introduced approximately 30 years ago in the south in the Parana state, primarily as a means of reducing erosion. Subsequently, research has begun to study the management of the crop waste products and their effects on soil fertility, either in terms of phosphorus management, as a means of controlling soil acidity, or determining how manures can be applied in a more localised manner. The spread of no-till in Brazil has involved a large amount of extension work. The area under no-tillage is still increasing in the centre and north of the country and currently occupies ca. 20 million hectares, covering a diversity of environmental conditions, cropping systems and management practices. Most studies of Brazilian soils give rates of carbon storage in the top 40 cm of the soil of 0.4 to 1.7 t C ha(-1) per year, with the highest rates in the Cerrado region. However, caution must be taken when analysing DMC systems in terms of carbon sequestration. Comparisons should include changes in trace gas fluxes and should not be limited to a consideration of carbon storage in the soil alone if the full implications for global warming are to be assessed.

Global vegetation and terrestrial carbon cycle changes after the last ice age

•In current models, the ecophysiological effects of CO2 create both woody thickening and terrestrial carbon uptake, as observed now, and forest cover and terrestrial carbon storage increases that took place after the last glacial maximum (LGM). Here, we aimed to assess the realism of modelled vegetation and carbon storage changes between LGM and the pre-industrial Holocene (PIH). •We applied Land Processes and eXchanges (LPX), a dynamic global vegetation model (DGVM), with lowered CO2 and LGM climate anomalies from the Palaeoclimate Modelling Intercomparison Project (PMIP II), and compared the model results with palaeodata. •Modelled global gross primary production was reduced by 27–36% and carbon storage by 550–694 Pg C compared with PIH. Comparable reductions have been estimated from stable isotopes. The modelled areal reduction of forests is broadly consistent with pollen records. Despite reduced productivity and biomass, tropical forests accounted for a greater proportion of modelled land carbon storage at LGM (28–32%) than at PIH (25%). •The agreement between palaeodata and model results for LGM is consistent with the hypothesis that the ecophysiological effects of CO2 influence tree–grass competition and vegetation productivity, and suggests that these effects are also at work today.

Direct soil moisture controls of future global soil carbon changes: An important source of uncertainty

The nature of the climate–carbon cycle feedback depends critically on the response of soil carbon to climate, including changes in moisture. However, soil moisture–carbon feedback responses have not been investigated thoroughly. Uncertainty in the response of soil carbon to soil moisture changes could arise from uncertainty in the relationship between soil moisture and heterotrophic respiration. We used twelve soil moisture–respiration functions (SMRFs) with a soil carbon model (RothC) and data from a coupled climate–carbon cycle general circulation model to investigate the impact of direct heterotrophic respiration dependence on soil moisture on the climate carbon cycle feedback. Global changes in soil moisture acted to oppose temperature‐driven decreases in soil carbon and hence tended to increase soil carbon storage. We found considerable uncertainty in soil carbon changes due to the response of soil respiration to soil moisture. The use of different SMRFs resulted in both large losses and small gains in future global soil carbon stocks, whether considering all climate forcings or only moisture changes. Regionally, the greatest range in soil carbon changes across SMRFs was found where the largest soil carbon changes occurred. Further research is needed to constrain the soil moisture–respiration relationship and thus reduce uncertainty in climate–carbon cycle feedbacks. There may also be considerable uncertainty in the regional responses of soil carbon to soil moisture changes since climate model predictions of regional soil moisture changes are less coherent than temperature changes.

The role of soil microbes in the global carbon cycle: tracking the below-ground microbial processing of plant-derived carbon for manipulating carbon dynamics in agricultural systems.

It is well known that atmospheric concentrations of carbon dioxide (CO2) (and other greenhouse gases) have increased markedly as a result of human activity since the industrial revolution. It is perhaps less appreciated that natural and managed soils are an important source and sink for atmospheric CO2 and that, primarily as a result of the activities of soil microorganisms, there is a soil-derived respiratory flux of CO2 to the atmosphere that overshadows by tenfold the annual CO2 flux from fossil fuel emissions. Therefore small changes in the soil carbon cycle could have large impacts on atmospheric CO2 concentrations. Here we discuss the role of soil microbes in the global carbon cycle and review the main methods that have been used to identify the microorganisms responsible for the processing of plant photosynthetic carbon inputs to soil. We discuss whether application of these techniques can provide the information required to underpin the management of agro-ecosystems for carbon sequestration and increased agricultural sustainability. We conclude that, although crucial in enabling the identification of plant-derived carbon-utilising microbes, current technologies lack the high-throughput ability to quantitatively apportion carbon use by phylogentic groups and its use efficiency and destination within the microbial metabolome. It is this information that is required to inform rational manipulation of the plant–soil system to favour organisms or physiologies most important for promoting soil carbon storage in agricultural soil.

Simulated last glacial maximum D14Catm and the deep glacial ocean carbon reservoir

∆14Catm has been estimated as 420 ± 80‰ (IntCal09) during the Last Glacial Maximum (LGM) compared to preindustrial times (0‰), but mechanisms explaining this difference are not yet resolved. ∆14Catm is a function of both cosmogenic production in the high atmosphere and of carbon cycling and partitioning in the Earth system. 10Be-based reconstructions show a contribution of the cosmogenic production term of only 200 ± 200‰ in the LGM. The remaining 220‰ have thus to be explained by changes in the carbon cycle. Recently, Bouttes et al. (2010, 2011) proposed to explain most of the difference in pCO2atm and δ13C between glacial and interglacial times as a result of brine-induced ocean stratification in the Southern Ocean. This mechanism involves the formation of very saline water masses that contribute to high carbon storage in the deep ocean. During glacial times, the sinking of brines is enhanced and more carbon is stored in the deep ocean, lowering pCO2atm. Moreover, the sinking of brines induces increased stratification in the Southern Ocean, which keeps the deep ocean well isolated from the surface. Such an isolated ocean reservoir would be characterized by a low ∆14C signature. Evidence of such 14C-depleted deep waters during the LGM has recently been found in the Southern Ocean (Skinner et al. 2010). The degassing of this carbon with low ∆14C would then reduce ∆14Catm throughout the deglaciation. We have further developed the CLIMBER-2 model to include a cosmogenic production of 14C as well as an interactive atmospheric 14C reservoir. We investigate the role of both the sinking of brine and cosmogenic production, alongside iron fertilization mechanisms, to explain changes in ∆14Catm during the last deglaciation. In our simulations, not only is the sinking of brine mechanism consistent with past ∆14C data, but it also explains most of the differences in pCO2atm and ∆14Catm between the LGM and preindustrial times. Finally, this study represents the first time to our knowledge that a model experiment explains glacial-interglacial differences in pCO2atm, δ13C, and ∆14C together with a coherent LGM climate.

Enhanced Australian carbon sink despite increased wildfire during the 21st century

Climate projections show Australia becoming significantly warmer during the 21st century, and precipitation decreasing over much of the continent. Such changes are conventionally considered to increase wildfire risk. Nevertheless, we show that burnt area increases in southern Australia, but decreases in northern Australia. Overall the projected increase in fire is small (0.72–1.31% of land area, depending on the climate scenario used), and does not cause a decrease in carbon storage. In fact, carbon storage increases by 3.7–5.6 Pg C (depending on the climate scenario used). Using a process-based model of vegetation dynamics, vegetation–fire interactions and carbon cycling, we show increased fire promotes a shift to more fire-adapted trees in wooded areas and their encroachment into grasslands, with an overall increase in forested area of 3.9–11.9%. Both changes increase carbon uptake and storage. The increase in woody vegetation increases the amount of coarse litter, which decays more slowly than fine litter hence leading to a relative reduction in overall heterotrophic respiration, further reducing carbon losses. Direct CO2 effects increase woody cover, water-use efficiency and productivity, such that carbon storage is increased by 8.5–14.8 Pg C compared to simulations in which CO2 is held constant at modern values. CO2 effects tend to increase burnt area, fire fluxes and therefore carbon losses in arid areas, but increase vegetation density and reduce burnt area in wooded areas.

The hidden organic carbon in deep mineral soils

Aims Current estimates of soil organic carbon (SOC) are based largely on surficial measurements to depths of 0.3 to 1 m. Many of the world’s soils greatly exceed 1 m depth and there are numerous reports of biological activity to depths of many metres. Although SOC storage to depths of up to 8 m has been previously reported, the extent to which SOC is stored at deeper depths in soil profiles is currently unknown. This paper aims to provide the first detailed analysis of these previously unreported stores of SOC. Methods Soils from five sites in the deeply weathered regolith in the Yilgarn Craton of south-western Australia were sampled and analysed for total organic carbon by combustion chromatography. These soils ranged between 5 and 38 m (mean 21 m) depth to bedrock and had been either recently reforested with Pinus pinaster or were under agriculture. Sites had a mean annual rainfall of between 399 and 583 mm yr−1. Results The mean SOC concentration across all sites was 2.30 ± 0.26 % (s.e.), 0.41 ± 0.05 % and 0.23 ± 0.04 % in the surface 0.1, 0.1–0.5 and 0.5 to 1.0 m increments, respectively. The mean value between 1 and 5 m was 0.12 ± 0.01 %, whereas between 5 and 35 m the values decreased from 0.04 ± 0.002 % to 0.03 ± 0.003 %. Mean SOC mass densities for each of the five locations varied from 21.8–37.5 kg C m−2, and were in toto two to five times greater than would be reported with sampling to a depth of 0.5 m. Conclusions This finding may have major implications for estimates of global carbon storage and modelling of the potential global impacts of climate change and land-use change on carbon cycles. The paper demonstrates the need for a reassessment of the current arbitrary shallow soil sampling depths for assessing carbon stocks, a revision of global SOC estimates and elucidation of the composition and fate of deep carbon in response to land use and climate change

Legacy effects of grassland management on soil carbon to depth

The importance of managing land to optimise carbon sequestration for climate change mitigation is widely recognised, with grasslands being identified as having the potential to sequester additional carbon. However, most soil carbon inventories only consider surface soils, and most large scale surveys group ecosystems into broad habitats without considering management intensity. Consequently, little is known about the quantity of deep soil carbon and its sensitivity to management. From a nationwide survey of grassland soils to 1 m depth, we show that carbon in grasslands soils is vulnerable to management and that these management effects can be detected to considerable depth down the soil profile, albeit at decreasing significance with depth. Carbon concentrations in soil decreased as management intensity increased, but greatest soil carbon stocks (accounting for bulk density differences), were at intermediate levels of management. Our study also highlights the considerable amounts of carbon in sub-surface soil below 30cm, which is missed by standard carbon inventories. We estimate grassland soil carbon in Great Britain to be 2097 Tg C to a depth of 1 m, with ~60% of this carbon being below 30cm. Total stocks of soil carbon (t ha-1) to 1 m depth were 10.7% greater at intermediate relative to intensive management, which equates to 10.1 t ha-1 in surface soils (0-30 cm), and 13.7 t ha-1 in soils from 30-100 cm depth. Our findings highlight the existence of substantial carbon stocks at depth in grassland soils that are sensitive to management. This is of high relevance globally, given the extent of land cover and large stocks of carbon held in temperate managed grasslands. Our findings have implications for the future management of grasslands for carbon storage and climate mitigation, and for global carbon models which do not currently account for changes in soil carbon to depth with management.

Climate change and the British Uplands: evidence for decision-making

We summarise the work of an interdisciplinary network set up to explore the impacts of climate change in the British Uplands. In this CR Special, the contributors present the state of knowledge and this introduction synthesises this knowledge and derives implications for decision makers. The Uplands are valued semi-natural habitats, providing ecosystem services that have historically been taken for granted. For example, peat soils, which are mostly found in the Uplands, contain around 50% of the terrestrial carbon in the UK. Land management continues to be a driver of ecosystem service delivery. Degraded and managed peatlands are subject to erosion and carbon loss with negative impacts on biodiversity, carbon storage and water quality. Climate change is already being experienced in British Uplands and is likely to exacerbate these pressures. Climate envelope models suggest as much as 50% of British Uplands and peatlands will be exposed to climate stress by the end of the 21st century under low and high emissions scenarios. However, process-based models of the response of organic soils to this climate stress do not give a consistent indication of what this will mean for soil carbon: results range from a very slight increase in uptake, through a clear decline, to a net carbon loss. Preserving existing peat stocks is an important climate mitigation strategy, even if new peat stops forming. Preserving upland vegetation cover is a key win–win management strategy that will reduce erosion and loss of soil carbon, and protect a variety of services such as the continued delivery of a high quality water resource.

Assessment of projected changes in upland environments using simple climatic indices

Climate controls upland habitats, soils and their associated ecosystem services; therefore, understanding possible changes in upland climatic conditions can provide a rapid assessment of climatic vulnerability over the next century. We used 3 different climatic indices that were optimised to fit the upland area classified by the EU as a Severely Disadvantaged Area (SDA) 1961–1990. Upland areas within the SDA covered all altitudinal ranges, whereas the maximum altitude of lowland areas outside of the SDA was ca. 300 m. In general, the climatic index based on the ratio between annual accumulated temperature (as a measure of growing season length) and annual precipitation predicted 96% of the SDA mapped area, which was slightly better than those indices based on annual or seasonal water deficit. Overall, all climatic indices showed that upland environments were exposed to some degree of change by 2071–2100 under UKCIP02 climate projections for high and low emissions scenarios. The projected area declined by 13 to 51% across 3 indices for the low emissions scenario and by 24 to 84% for the high emissions scenario. Mean altitude of the upland area increased by +11 to +86 m for the low scenario and +21 to +178 m for the high scenario. Low altitude areas in eastern and southern Great Britain were most vulnerable to change. These projected climatic changes are likely to affect upland habitat composition, long-term soil carbon storage and wider ecosystem service provision, although it is not yet possible to determine the rate at which this might occur.