Comparative genomics of Mortierella elongata and its bacterial endosymbiont Mycoavidus cysteinexigens.

Endosymbiosis of bacteria by eukaryotes is a defining feature of cellular evolution. In addition to well-known bacterial origins for mitochondria and chloroplasts, multiple origins of bacterial endosymbiosis are known within the cells of diverse animals, plants and fungi. Early-diverging lineages of terrestrial fungi harbor endosymbiotic bacteria belonging to the Burkholderiaceae. We sequenced the metagenome of the soil-inhabiting fungus Mortierella elongata and assembled the complete circular chromosome of its endosymbiont, Mycoavidus cysteinexigens, which we place within a lineage of endofungal symbionts that are sister clade to Burkholderia. The genome of M. elongata strain AG77 features a core set of primary metabolic pathways for degradation of simple carbohydrates and lipid biosynthesis, while the M. cysteinexigens (AG77) genome is reduced in size and function. Experiments using antibiotics to cure the endobacterium from the host demonstrate that the fungal host metabolism is highly modulated by presence/absence of M. cysteinexigens. Independent comparative phylogenomic analyses of fungal and bacterial genomes are consistent with an ancient origin for M. elongata - M. cysteinexigens symbiosis, most likely over 350 million years ago and concomitant with the terrestrialization of Earth and diversification of land fungi and plants.

Plant water relations at elevated CO2 -- implications for water-limited environments.

Long-term exposure of plants to elevated [CO2] leads to a number of growth and physiological effects, many of which are interpreted in the context of ameliorating the negative impacts of drought. However, despite considerable study, a clear picture in terms of the influence of elevated [CO2] on plant water relations and the role that these effects play in determining the response of plants to elevated [CO2] under water-limited conditions has been slow to emerge. In this paper, four areas of research are examined that represent critical, yet uncertain, themes related to the response of plants to elevated [CO2] and drought. These include (1) fine-root proliferation and implications for whole-plant water uptake; (2) enhanced water-use efficiency and consequences for drought tolerance; (3) reductions in stomatal conductance and impacts on leaf water potential; and (4) solute accumulation, osmotic adjustment and dehydration tolerance of leaves. A survey of the literature indicates that the growth of plants at elevated [CO2] can lead to conditions whereby plants maintain higher (less negative) leaf water potentials. The mechanisms that contribute to this effect are not fully known, although CO2-induced reductions in stomatal conductance, increases in whole-plant hydraulic conductance and osmotic adjustment may be important. Less understood are the interactive effects of elevated [CO2] and drought on fine-root production and water-use efficiency, and the contribution of these processes to plant growth in water-limited environments. Increases in water-use efficiency and reductions in water use can contribute to enhanced soil water content under elevated [CO2]. Herbaceous crops and grasslands are most responsive in this regard. The conservation of soil water at elevated [CO2] in other systems has been less studied, but in terms of maintaining growth or carbon gain during drought, the benefits of CO2-induced improvements in soil water content appear relatively minor. Nonetheless, because even small effects of elevated [CO2] on plant and soil water relations can have important implications for ecosystems, we conclude that this area of research deserves continued investigation. Future studies that focus on cellular mechanisms of plant response to elevated [CO2] and drought are needed, as are whole-plant investigations that emphasize the integration of processes throughout the soil--plant--atmosphere continuum. We suggest that the hydraulic principles that govern water transport provide an integrating framework that would allow CO2-induced changes in stomatal conductance, leaf water potential, root growth and other processes to be uniquely evaluated within the context of whole-plant hydraulic conductance and water transport efficiency.

Growth and carbohydrate status of coppice shoots of hybrid poplar following shoot pruning.

Fifteen, 1-year-old Populus maximowiczii Henry x P. nigra L. 'MN9' trees were decapitated and allowed to sprout. After 8 weeks, all had 6 to 10 coppice shoots. All shoots, except the tallest (dominant) shoot, were removed from five of the trees (pruned treatment), and shoot growth, gas exchange and carbohydrate status were compared in the pruned and unpruned trees. Although photosynthetic rate of recently mature leaves of pruned trees was approximately 50% greater than that of leaves on the dominant shoot of unpruned trees, and the dry weight of leaves of pruned trees was 37% greater than that of the leaves on the dominant shoot of unpruned trees, the shoot dry matter relative growth rate did not differ between treatments. Concentrations of water-soluble carbohydrates and starch in the uppper stem and leaves of the dominant shoot were similar in pruned and unpruned trees. However, relative to that of the dominant shoot in unpruned trees, the lower stem in pruned trees was depleted in both soluble carbohydrates and starch. Starch deposition, assessed as the quantity of (14)C-starch in tissues 24 h after a fully expanded source leaf was labeled with (14)CO(2), was 3.9 times greater in roots of pruned trees than in roots of unpruned trees. We conclude that early removal of all but the dominant shoot reduces the carbohydrate status of the roots and the lower portion of the stem by eliminating the excised shoots as a source of photosynthate.

Interactions between drought and elevated CO2 on growth and gas exchange of seedlings of three deciduous tree species.

Interactions between elevated atmospheric CO2 and drought on growth and gas exchange of American sycamore (Platanus occidentalis L.), sweetgum (Liquidambar styraciflua L.) and sugar maple (Acer saccharum Marsh.) were investigated using 1-yr-old seedlings, planted in 8 1 pots and grown in four open-top chambers, containing either ambient air or ambient air enriched with 300 µmol mol-1 CO2 . Two soil moisture regimes were included within each chamber: a 'well-watered' treatment with plants watered daily and a 'drought' treatment in which plants were subjected to a series of drought cycles. Duration and depth of the drought cycles were determined by soil matric potential. Mean soil water potential at rewatering for the water-stressed seedlings under ambient CO2 for sugar maple, sweetgum and sycamore was -0.5, -0.7 and -1.8 MPa, respectively, compared with > -0.1 MPa for all well-watered plants. Elevated CO2 increased relative growth rate of well-watered sugar maple by 181%, resulting in a 4.3-fold increase in total plant dry weight after 81 d, compared with 1.4 and 1.6-fold increases for sweetgum and sycamore, respectively, after 69 d. Although elevated CO2 increased net CO2 assimilation rate of sugar maple by 115%, there was a 10-fold increase in leaf area which contributed to the growth response. Although drought did not eliminate a growth response of sugar maple to elevated CO2 it greatly reduced the elevated CO2 -induced enhancement of relative growth rate. In contrast, relative growth rates of sweetgum and sycamore were not significantly increased by elevated CO2 . Drought, under elevated CO2 , reduced leaf area of all three species to a greater extent than it reduced net CO2 assimilation rate. The response ranged from no effect in sugar maple to a 40 % reduction in sycamore, with sweetgum exhibiting an intermediate response. Results indicate that drought may alter the growth response, gas exchange and water relations of tree species growing in an elevated CO2 atmosphere. Under high nutrient and water availability, sugar maple may benefit the most (of the three species studied) from a CO2 - enriched atmosphere, but productivity gains will be limited if frequent drought is prevalent.

Carbohydrate mobilization following shoot defoliation and decapitation in hybrid poplar.

The effects of shoot defoliation, decapitation, and disbudding on carbon mobilization were investigated in rooted cuttings of Populus maximowiczii x nigra L. 'MN9'. Ten days after complete shoot defoliation or decapitation, the stem starch concentration of treated plants declined to one-half that of intact plants, and there were similar or greater reductions in the concentrations of glucose, fructose, sucrose, galactose, and shikimic acid. Partial shoot defoliation (50%) and complete disbudding had no effect on stem starch concentration, but stem sucrose concentration was reduced in all treatments. Sucrose depletion preceded and may have induced other changes in the carbon status of plants subjected to leaf or shoot removal. Four days after shoot decapitation, the sucrose concentration of roots of treated plants was reduced to 25% of that of intact plants. However, the concentrations of fructose and glucose increased in the roots of treated plants and was followed by the accumulation of shikimic acid, salicyl alcohol, unknown compound A and salicin. The possible role of increased concentrations of root organic solutes in the water relations and regrowth process of decapitated plants is discussed.

Responses of loblolly pine seedlings to elevated CO(2) and fluctuating water supply.

Osmotic adjustment of loblolly pine (Pinus taeda L.) seedlings to fluctuating water supply in elevated CO(2) was investigated. Seedlings were grown in controlled-environment chambers in either 350 or 700 micro l l(-1) CO(2) with weekly watering for four months, after which they were either watered weekly (well-watered treatment) or every two weeks (water-stress treatment) for 59 days. Osmotic adjustment was assessed by pressure-volume analysis of shoots and by analysis of soluble carbohydrates and free amino acids in roots during the last drying cycle. In well-watered seedlings, elevated CO(2) increased the concentration of soluble sugars in roots by 68%. Water stress reduced the soluble sugar concentration in roots of seedling growing in ambient CO(2) to 26% of that in roots of well-watered seedlings. Elevated CO(2) mitigated the water stress-induced decrease in the concentration of soluble sugars in roots. However, this was probably due, in part, to carbohydrate loading during the first four months when all seedlings were grown in the presence of a high water supply, rather than to osmotic adjustment to water stress. Water stress caused a doubling in the concentration of free primary amino acids in roots, whereas elevated CO(2) reduced primary amino acid and nitrogen concentrations to 32 and 74%, respectively, of those in roots of seedlings grown in ambient CO(2). There was no indication of large-scale osmotic adjustment to water stress or that elevated CO(2) enhanced osmotic adjustment in loblolly pine.