Gretchen B. North

Plasma Membrane Aquaporins Play a Significant Role during Recovery from Water Deficit

The role of plasma membrane aquaporins (PIPs) in water relations of Arabidopsis was studied by examining plants with reduced expression of PIP1 and PIP2 aquaporins, produced by crossing two different antisense lines. Compared with controls, the double antisense (dAS) plants had reduced amounts of PIP1 and PIP2 aquaporins, and the osmotic hydraulic conductivity of isolated root and leaf protoplasts was reduced 5- to 30-fold. The dAS plants had a 3-fold decrease in the root hydraulic conductivity expressed on a root dry mass basis, but a compensating 2.5-fold increase in the root to leaf dry mass ratio. The leaf hydraulic conductance expressed on a leaf area basis was similar for the dAS compared with the control plants. As a result, the hydraulic conductance of the whole plant was unchanged. Under sufficient and under water-deficient conditions, stomatal conductance, transpiration rate, plant hydraulic conductance, leaf water potential, osmotic pressure, and turgor pressure were similar for the dAS compared with the control plants. However, after 4 d of rewatering following 8 d of drying, the control plants recovered their hydraulic conductance and their transpiration rates faster than the dAS plants. Moreover, after rewatering, the leaf water potential was significantly higher for the control than for the dAS plants. From these results, we conclude that the PIPs play an important role in the recovery of Arabidopsis from the water-deficient condition.

Plant compartment and biogeography affect microbiome composition in cultivated and native Agave species

Summary Desert plants are hypothesized to survive the environmental stress inherent to these regions in part thanks to symbioses with microorganisms, and yet these microbial species, the communities they form, and the forces that influence them are poorly understood. Here we report the first comprehensive investigation of the microbial communities associated with species of Agave, which are native to semiarid and arid regions of Central and North America and are emerging as biofuel feedstocks. We examined prokaryotic and fungal communities in the rhizosphere, phyllosphere, leaf and root endosphere, as well as proximal and distal soil samples from cultivated and native agaves, through Illumina amplicon sequencing. Phylogenetic profiling revealed that the composition of prokaryotic communities was primarily determined by the plant compartment, whereas the composition of fungal communities was mainly influenced by the biogeography of the host species. Cultivated A. tequilana exhibited lower levels of prokaryotic diversity compared with native agaves, although no differences in microbial diversity were found in the endosphere. Agaves shared core prokaryotic and fungal taxa known to promote plant growth and confer tolerance to abiotic stress, which suggests common principles underpinning Agave–microbe interactions.

Drought-induced changes in soil contact and hydraulic conductivity for roots of Opuntia ficus-indica with and without rhizosheaths

Water movement between roots and soil can be limited by incomplete root–soil contact, such as that caused by air gaps due to root shrinkage, and can also be influenced by rhizosheaths, composed of soil particles bound together by root exudates and root hairs. The possible occurrence of air gaps between the roots and the soil and their consequences for the hydraulic conductivity of the root–soil pathway were therefore investigated for the cactus t Opuntia ficus-indica, which has two distinct root regions: a younger, distal region where rhizosheaths occur, and an older, proximal region where roots are bare. Resin-embedded sections of roots in soil were examined microscopically to determine root–soil contact for container-grown plants kept moist for 21 days, kept moist and vibrated to eliminate air gaps, droughted for 21 days, or droughted and vibrated. During drought, roots shrank radially by 30% and root–soil contact in the bare root region of nonvibrated containers was reduced from 81% to 31%. For the sheathed region, the hydraulic conductivity of the rhizosheath was the least limiting factor and the root hydraulic conductivity was the most limiting; for the bare root region, the hydraulic conductivity of the soil was the least limiting factor and the hydraulic conductivity of the root–soil air gap was the most limiting. The rhizosheath, by virtually eliminating root–soil air gaps, facilitated water uptake in moist soil. In the bare root region, the extremely low hydraulic conductivity of the root–soil air gap during drought helped limit water loss from roots to a drier soil.

Soil Sheaths, Photosynthate Distribution to Roots, and Rhizosphere Water Relations for Opuntia ficus-indica

Soil sheaths incorporating aggregated soil particles surround young roots of many species, but the effects of such sheaths on water movement between roots and the soil are largely unknown. The quantity and location of root exudates associated with soil sheath formation and root water loss were therefore examined for Opuntia ficus-indica, which has prominent soil sheaths along the entire length of its young roots, except within 1.4 cm of the tip. The soil sheaths, which averaged 0.7 mm in thickness, were composed of soil particles and root hairs, both of which were covered with exuded mucilaginous material. As determined with a 14C pulse-labeling technique, 2% of newly fixed 14C-photosynthate was translocated into the roots at 3 d, 6% at 9 d, and 8% at 15 d after labeling. The fraction of insoluble 14C in the roots increased twofold from 3 d to 15 d. Over the same time period, 6%-9% of the 14C translocated to the roots was exuded into the soil. The soluble 14C compounds exuded into the soil were greater in the 3-cm segment at the root tip than elsewhere along the root, whereas mucilage was exuded relatively uniformly along roots 15 cm in length. The volumetric efflux of water increased for both sheathed and unsheathed roots as the soil water potential decreased from -0.1 MPa to -1.0 MPa. The efflux rate was greater for unsheathed roots than for sheathed roots, which were more turgid and had a higher water potential, especially at lower soil water potentials. During drying, soil particles in the sheaths aggregate more tightly, making the sheaths less permeable to water and possibly creating air gaps. The soil sheaths of O. ficus-indica thus reduce water loss from the roots to a drying soil.

Leaf hydraulic conductance for a tank bromeliad: axial and radial pathways for moving and conserving water.

Epiphytic plants in the Bromeliaceae known as tank bromeliads essentially lack stems and absorptive roots and instead take up water from reservoirs formed by their overlapping leaf bases. For such plants, leaf hydraulic conductance is plant hydraulic conductance. Their simple strap-shaped leaves and parallel venation make them suitable for modeling leaf hydraulic conductance based on vasculature and other anatomical and morphological traits. Plants of the tank bromeliad Guzmania lingulata were investigated in a lowland tropical forest in Costa Rica and a shaded glasshouse in Los Angeles, CA, USA. Stomatal conductance to water vapor and leaf anatomical variables related to hydraulic conductance were measured for both groups. Tracheid diameters and numbers of vascular bundles (veins) were used with the Hagen–Poiseuille equation to calculate axial hydraulic conductance. Measurements of leaf hydraulic conductance using the evaporative flux method were also made for glasshouse plants. Values for axial conductance and leaf hydraulic conductance were used in a model based on leaky cable theory to estimate the conductance of the radial pathway from the vein to the leaf surface and to assess the relative contributions of both axial and radial pathways. In keeping with low stomatal conductance, low stomatal density, low vein density, and narrow tracheid diameters, leaf hydraulic conductance for G. lingulata was quite low in comparison with most other angiosperms. Using the predicted axial conductance in the leaky cable model, the radial resistance across the leaf mesophyll was predicted to predominate; lower, more realistic values of axial conductance resulted in predicted radial resistances that were closer to axial resistance in their impact on total leaf resistance. Tracer dyes suggested that water uptake through the tank region of the leaf was not limiting. Both dye movement and the leaky cable model indicated that the leaf blade of G. lingulata was structurally and hydraulically well-suited to conserve water.

Plant hydraulics: new discoveries in the pipeline

Increasing numbers of plant scientists are recognizing the importance of hydraulic design in determining plant function. Hydraulic design – which can be broadly defined as the functional properties of the plant vascular system – is a determinant not only of plant water balance but also of photosynthetic rates and ecological niche differentiation. Classic approaches (Tyree & Zimmermann, 2002) and newer concepts (Holbrook & Zwieniecki, 2005) are being applied to questions central to the evolution and ecology of plant species, ranging from organ to organism to ecosystem. A recent workshop held in southern California reflected diverse research programs but also highlighted a convergence of interest on key questions and promising approaches. Several breakout sessions focused on defining pressing questions of plant hydraulics and on addressing the critical need for standardization of practices for research on these topics.

Root deployment and shoot growth for two desert species in response to soil rockiness

Soil texture, as well as the presence of rocks, can determine the water status, growth, and distribution of plants in arid environments. The effects of soil rockiness and soil particle size distribution on shoot and root growth, root system size, rooting depth, and water relations were therefore investigated for the Crassulacean acid metabolism leaf succulent Agave deserti and the C(4) bunchgrass Pleuraphis rigida after precipitation events during the summer and winter/spring rainfall periods in the northwestern Sonoran Desert. The soils at the field site varied from sandy (<3% rocks by volume) to rocky (up to 35% rocks), with greater water availability at higher water potentials for sandy than for rocky soils. Although A. deserti was absent from the sandiest sites, its shoot and root growth during both rainfall periods were greatest in comparatively sandier sites and decreased as the soil rock content increased. Furthermore, A. deserti had twofold greater root surface area, root : leaf area ratio, and mean rooting depth at sandier than at rocky sites. As for A. deserti, shoot growth was greater for P. rigida at the sandier sites than at the rockier sites, even though its root surface area and mean rooting depth did not vary significantly. After early spring rainfall events, the leaf water potential for A. deserti did not differ between rocky and sandy sites, but transpiration rates were almost twofold greater at rocky than at sandy sites. During the same period, P. rigida had lower leaf water potentials and 25% lower transpiration rates at rocky than at sandy sites. The greater variability in the deployment of the root systems of A. deserti in response to soil rockiness may reflect its evergreen habit and slower growth, which allow it to endure periods of lower water availability than does P. rigida, whose leaves die during drought.

Allocation tradeoffs among chaparral shrub seedlings with different life history types (Rhamnaceae)

PREMISE OF THE STUDY California chaparral shrub species have different life history types: Nonsprouters (NS) are killed by fire and persist through a fire-stimulated seed bank; facultative sprouters (FS) reestablish by a combination of vegetative sprouting and seeding; and obligate sprouters (OS) reestablish exclusively by sprouting. Nonsprouters and FS establish seedlings in open-canopy postfire environments, whereas OS establish seedlings between fires in the shady understory. We hypothesized that allocation differences among seedlings of postfire sprouters and nonsprouters and regeneration niche differences would lead to contrasting patterns in biomass accumulation (NS > FS > OS, in sun; OS > FS > NS, in shade). METHODS Seedlings of three species from each life history type were grown in sun and 75% shade. We measured net carbon assimilation and biomass accumulation after one year. KEY RESULTS Biomass accumulation was similar in the sun except FS>OS. In the shade, NS had lower biomass than FS and OS. Assimilation rates, nitrogen relations, and allocation differences could not fully explain biomass accumulation differences. Instead, biomass accumulation was inversely related to water-stress tolerance and shade tolerance. Additionally, OS and FS differed in root/shoot allocation even though both are sprouters. CONCLUSIONS Seedling growth and carbon assimilation rates were divergent among three life history types and were consistent with differences in tolerance to water stress and shade or sun regeneration niches, but not tradeoffs in sprouting-related allocation differences per se.

Water flow in roots: structural and regulatory features

Publisher Summary This chapter examines the structural components of the radial pathway for water flow in roots and discusses possible regulators of water movement through both radial and axial pathways, specifically, aquaporins and embolism, respectively, and how these change during development and water stress. It considers how water stress might also affect the proportional limitations imposed by the two pathways on overall water transport by roots. In herbaceous plants in moist soil, rates of axial flow are usually high enough that the primary limitation for water transport occurs in the radial pathway (i.e., from the ambient solution into the root xylem). The details of radial water flow are less well understood than those of axial flow, however, because roots are highly complex, and the proportions of water moving in the various available pathways may be altered depending on the driving forces. The anatomy of roots changes along their length as various structures develop, mature, and, perhaps, die. Their structure can change in response to different environmental conditions. In addition, there is wide variation in structure amongst species. Despite the complexities of roots and the challenges involved in determining how water moves through them, recent progress has been made on several fronts.

Contractile roots in succulent monocots: convergence, divergence and adaptation to limited rainfall

Contractile roots (CRs) that pull shoots further down in the soil are a possible example of convergent evolution in two monocot families, the Agavaceae and the Asphodelaceae. The association between CRs, water uptake and habitat aridity was investigated for agaves, yuccas and aloes by assessing the occurrence of CRs and the amount of root contraction for glasshouse-grown plants with respect to mean annual rainfall of their native habitats. Structural features of CRs as well as root hydraulic conductance were compared with those of non-contractile roots (NCRs). CRs occurred in 55% of the 73 species examined, including 64% of the agaves and 85% of the yuccas, but in none of the aloes despite the occurrence of CRs in related genera. The phylogenetic distribution of CRs was consistent with multiple acquisitions or losses of the trait. The amount of root contraction showed a highly significant negative relationship with mean annual rainfall, although other environmental factors may also be important. Radial hydraulic conductance of the basal (contractile) zone exceeded that of the midroot zone for CRs; for NCRs, the opposite was true. Thus, CRs in the species examined may provide a mechanism for greater water uptake near the soil surface in regions with limited rainfall.