Fire and herbivory drive fungal and bacterial communities through distinct above- and belowground mechanisms.

Fire and herbivory are important natural disturbances in grassy biomes. Both drivers are likely to influence belowground microbial communities but no studies have unravelled the long-term impact of both fire and herbivory on bacterial and fungal communities. We hypothesized that soil bacterial communities change through disturbance-induced shifts in soil properties (e.g. pH, nutrients) while soil fungal communities change through vegetation modification (biomass and species composition). To test these ideas, we characterised soil physico-chemical properties (pH, acidity, C, N, P and exchangeable cations content, texture, bulk density, moisture), plant species richness and biomass, microbial biomass and bacterial and fungal community composition and diversity (using 16S and ITS rRNA amplicon sequencing, respectively) in six long-term (18 to 70 years) ecological research sites in South African savanna and grassland ecosystems. We found that fire and herbivory regimes profoundly modified soil physico-chemical properties, plant species richness and standing biomass. In all sites, an increase in woody biomass (ranging from 12 to 50%) was observed when natural disturbances were excluded. The intensity and direction of changes in soil properties were highly dependent on the topo-pedo-climatic context. Overall, fire and herbivory shaped bacterial and fungal communities through distinct driving forces: edaphic properties (including Mg, pH, Ca) for bacteria, and vegetation (herbaceous biomass and woody cover) for fungi. Fire and herbivory explained on average 7.5 and 9.8% of the fungal community variability, respectively, compared to 6.0 and 5.6% for bacteria. The relatively small changes in microbial communities due to natural disturbance is in stark contrast to dramatic vegetation and edaphic changes and suggests that soil microbial communities, having evolved with disturbance, are resistant to change. This represents both a buffer to short-term anthropogenic-induced changes and a restoration challenge in the face of long-term changes.

Drivers of woody plant encroachment over Africa.

While global deforestation induced by human land use has been quantified, the drivers and extent of simultaneous woody plant encroachment (WPE) into open areas are only regionally known. WPE has important consequences for ecosystem functioning, global carbon balances and human economies. Here we report, using high-resolution satellite imagery, that woody vegetation cover over sub-Saharan Africa increased by 8% over the past three decades and that a diversity of drivers, other than CO2, were able to explain 78% of the spatial variation in this trend. A decline in burned area along with warmer, wetter climates drove WPE, although this has been mitigated in areas with high population growth rates, and high and low extremes of herbivory, specifically browsers. These results confirm global greening trends, thereby bringing into question widely held theories about declining terrestrial carbon balances and desert expansion. Importantly, while global drivers such as climate and CO2 may enhance the risk of WPE, managing fire and herbivory at the local scale provides tools to mitigate continental WPE.

Benefits of photosynthesis for insects in galls.

Insect-induced plant galls are predominantly reputed to act as strong carbon sinks, although many types of galls contain chlorophyll and have the potential to photosynthesize. We investigated whether the photosynthetic capacity of bud galls induced by a Pteromalid wasp, Trichilogaster acaciaelongifoliae, in Acacia longifolia subsidises carbon budgets or provides O(2) to the larvae while concurrently consuming CO(2) in the dense gall tissue, thereby maintaining (O(2)) and (CO(2)) within the range of larval tolerance. Low (O(2)) (<5 % v/v) were found within the internal tissues of galls, and these concentrations responded only marginally to light, suggesting that the photosynthetic activity within the gall is inconsequential in the provision of O(2) to the larvae. The metabolic response of larvae to reduced (O(2)) and elevated (CO(2)) indicated that larvae were tolerant of hypoxia/hypercarbia and also capable of reducing their respiratory rates to cope with hypercarbia. The low mortality of larvae in galls shaded with Al-foil for 20 days showed that photosynthesis was not vital for the survival of the larvae, although growth of shaded galls was substantially reduced. Gas exchange measurements confirmed that, while photosynthesis never fully compensated for the respiratory costs of galls, it contributed substantially to the maintenance and growth, especially of young galls, reducing their impact as carbon sinks on the host. We conclude that, although photosynthesis may contribute to O(2) provision, its main role is to reduce the dependence of the insect-induced gall on the host plant for photosynthates, thereby reducing intra-plant, inter-gall competition and enhancing the probability that each gall will reach maturity.

Supply and demand: sink regulation of sugar accumulation in sugarcane.

Sugarcane (Saccharum spp. hybrids) accumulates sucrose to high concentrations and, as a result, has been the focus of extensive research into the biochemistry and physiology of sucrose accumulation. Despite this, the relationship between source leaf photosynthetic activity and sucrose accumulation in the culm sink is not well understood. The observations that photosynthetic activity declines during culm maturation in commercial cultivars and that high-sucrose-accumulating noble ancestral genotypes (Saccharum officinarum L.) photosynthesize at rates two-thirds of those of low-sucrose ancestors (Saccharum spontaneum L.) indicate that source-sink communication may play a pivotal role in determining sucrose yield. Although maturation of the culm results in a decreased demand for sucrose, recent evidence from partial leaf shading, defoliation, and transgenic studies indicates that sugarcane cultivars are capable of further increases in sugar content. Furthermore, sugarcane leaves appear to retain the capacity to increase the supply of assimilate to culm tissues under conditions of increased assimilate demand. The relationship between source and sink tissues in sugarcane should be viewed within a supply-demand paradigm; an often neglected conceptual approach in the study of this crop. Uncoupling of the signalling pathways that mediate negative feedback between source and sink tissues may result in improved leaf assimilation rates and, consequently, lead to increased sugarcane sucrose yields.

Changes in photosynthetic rates and gene expression of leaves during a source-sink perturbation in sugarcane.

BACKGROUND AND AIMS: In crops other than sugarcane there is good evidence that the size and activity of carbon sinks influence source activity via sugar-related regulation of the enzymes of photosynthesis, an effect that is partly mediated through coarse regulation of gene expression. METHODS: In the current study, leaf shading treatments were used to perturb the source-sink balance in 12-month-old Saccharum spp. hybrid 'N19' (N19) by restricting source activity to a single mature leaf. Changes in leaf photosynthetic gas exchange variables and leaf and culm sugar concentrations were subsequently measured over a 14 d period. In addition, the changes in leaf gene response to the source-sink perturbation were measured by reverse northern hybridization analysis of an array of 128 expressed sequence tags (ESTs) related to photosynthetic and carbohydrate metabolism. KEY RESULTS: Sucrose concentrations in immature culm tissue declined significantly over the duration of the shading treatment, while a 57 and 88% increase in the assimilation rate (A) and electron transport rate (ETR), respectively, was observed in the source leaf. Several genes (27) in the leaf displayed a >2-fold change in expression level, including the upregulation of several genes associated with C(4) photosynthesis, mitochondrial metabolism and sugar transport. Changes in gene expression levels of several genes, including Rubisco (EC 4.1.1.39) and hexokinase (HXK; EC 2.7.1.1), correlated with changes in photosynthesis and tissue sugar concentrations that occurred subsequent to the source-sink perturbation. CONCLUSIONS: These results are consistent with the notion that sink demand may limit source activity through a kinase-mediated sugar signalling mechanism that correlates to a decrease in source hexose concentrations, which, in turn, correlate with increased expression of genes involved in photosynthesis and metabolite transport. The signal feedback system reporting sink sufficiency and regulating source activity may be a potentially valuable target for future genetic manipulation to increase sugarcane sucrose yield.

Does the prostrate-leaved geophyte Brunsvigia orientalis utilize soil-derived CO2 for photosynthesis?

BACKGROUND AND AIMS: A test was made of the hypothesis that the prostrate growth habit of the leaves of the geophyte Brunsvigia orientalis enables utilization of soil-derived CO(2) and is related to the presence of lysigenous air-filled channels characteristic of B. orientalis leaves. METHODS: Brunsvigia orientalis was sampled at a field site. Leaf anatomy, stomatal density, leaf/soil gas exchange characteristics and soil atmosphere and leaf delta(13)C isotope abundances were examined. KEY RESULTS: The leaves of B. orientalis have large lysigenous air-filled channels separating the upper and lower surfaces of the leaves. The upper surface comprised approx. 70 % of the leaf mass and 75 % of the leaf N (mmol g(-1)). Between 20 % and 30 % of the stomatal conductance and CO(2) assimilation was through the lower surface of the leaf. CO(2) efflux rates from the soil surface were up to 5.4 micromol m(-2) s(-1) while photosynthetic fluxes through the lower surface of the leaves were approx. 7 micromol m(-2) s(-1). However, the utilization of soil-derived CO(2) only altered the leaf delta(13)C isotope abundance of the prostrate leaves by a small amount. Using delta(13)C values it was estimated that 7 % of the leaf tissue C was derived from soil-derived CO(2). CONCLUSIONS: A small proportion of photosynthetically fixed CO(2) was derived from the soil, with minimal associated transpirational H(2)O loss into the space between the leaf and soil. The soil-derived CO(2), taken up through the lower surface was probably assimilated by the palisade tissue in the upper surface of the leaf which was exposed to sunlight and where most of the leaf N was located. The occurrence of lysigenous air channels in the leaves may provide longitudinal strength without impaired transfer of CO(2) taken up through the lower surface to the upper surface.

Sink strength regulates photosynthesis in sugarcane.

The relationship in sugarcane (Saccharum spp.) between photosynthetic source tissue and sink material was examined through manipulation of the sink:source ratio of field-grown Saccharum spp. hybrid cv. N19 (N19). To enhance sink strength, all leaves, except for the third fully expanded leaf, were enclosed in 90% shade cloth for varying periods of time. Variations in sucrose, glucose and fructose concentrations were measured and the effects of shading on the leaf gas exchange and fluorescence characteristics recorded. Changes in carbon partitioning caused by shading were examined based on the uptake and translocation of fixed 14CO2. Following a decline in sucrose concentrations in young internodal tissue and shaded leaves, significant increases in the CO2-saturated photosynthetic rate (Jmax), carboxylation efficiency (CE) and electron transport rate were observed in unshaded leaves after 8 d of shading treatment. It was concluded that up-regulation of source-leaf photosynthetic capacity is correlated with a decrease in assimilate availability to acropetal culm sink tissue. Furthermore, a significant relationship was revealed between source hexose concentration and photosynthetic activity.

The influence of root assimilated inorganic carbon on nitrogen acquisition/assimilation and carbon partitioning.

Understanding of the influences of root-zone CO2 concentration on nitrogen (N) metabolism is limited. The influences of root-zone CO2 concentration on growth, N uptake, N metabolism and the partitioning of root assimilated 14C were determined in tomato (Lycopersicon esculentum). Root, but not leaf, nitrate reductase activity was increased in plants supplied with increased root-zone CO2. Root phosphoenolpyruvate carboxylase activity was lower with NO3(-)- than with NH4(+)-nutrition, and in the latter, was also suppressed by increased root-zone CO2. Increased growth rate in NO3(-)-fed plants with elevated root-zone CO2 concentrations was associated with transfer of root-derived organic acids to the shoot and conversion to carbohydrates. With NH4(+)-fed plants, growth and total N were not altered by elevated root-zone CO2 concentrations, although 14C partitioning to amino acid synthesis was increased. Effects of root-zone CO2 concentration on N uptake and metabolism over longer periods (> 1 d) were probably limited by feedback inhibition. Root-derived organic acids contributed to the carbon budget of the leaves through decarboxylation of the organic acids and photosynthetic refixation of released CO2.

Variation in root-zone CO2 concentration modifies isotopic fractionation of carbon and nitrogen in tomato seedlings.

• The contribution to the carbon budget and growth by root acquisition of inorganic carbon and the influence that this has on NO 3 - and NH 4 + uptake and assimilation has not been adequately quantified. • The influence of varying root-zone CO 2 concentrations on tissue δ 13 C and δ 15 N was used to estimate the contribution to the carbon budget of root-assimilated carbon in tomato ( Lycopersicon esculentum ) seedlings. • Biomass accumulation was greater at 0.5% and 1% (v/v) root-zone CO 2 in NO 3 - and NH 4 + -fed plants than with 0% root-zone CO 2 . The plant δ 13 C values were not altered by 1% CO 2 with δ 13 C = -29.00‰, but they were increased when supplied with 1% CO 2 with δ 13 C = -10.91‰. The δ 15 N values of NO 3 - -fed plants were unchanged by variation in root-zone CO 2 concentration. In NH 4 + -fed plants the δ 15 N values were c.  1.5‰ higher at 1% CO 2 . • Changes in δ 13 C values with increased CO 2 concentration (δ 13 C = -10.91‰) were ascribed to root incorporation of CO 2 . Less than 5% of carbon was derived from root dark fixation and thus cannot explain increases in growth on a mass basis. Reduced discrimination with NH 4 + nutrition at 1% CO 2 could be related to increased exudation of NH 4 + and organic nitrogen and also reduced uptake.

Elevated root zone dissolved inorganic carbon can ameliorate aluminium toxicity in tomato seedlings.

• The potential amelioration of Al (Al) toxicity by elevated dissolved inorganic carbon (DIC = CO2  + HCO3 - ) in the root medium was investigated in both NH4 + - and NO3 - -fed Lycopersicon esculentum (tomato) plants. • Hydroponically grown L. esculentum seedlings were intermittently supplied with 50 µM AlCl3 and the nutrient solutions aerated with either 360 or 5000 ppm CO2 . Relative growth rate (RGR), nitrogen uptake, root respiration and root incorporation of DI14 C and subsequent partitioning were measured. • Al reduced the RGR of plants grown with 360 ppm root-zone CO2 . At elevated root-zone CO2 , Al had no significant effect on the RGR of NO3 - -fed plants whereas the RGR of the NH4 + -fed plants was increased by 21%. Al decreased the respiratory quotient (Rq) by 15% at 360 ppm CO2 , but had no influence at 2000 ppm CO2 . Exudation of organic 14 C, especially of [14 C]-organic acids derived from root incorporation of DI14 C, was increased by Al. • It is concluded that elevated DIC partially ameliorated Al toxicity by providing anaplerotic carbon for organic acid synthesis.