Does acclimation in cavitation resistance due to mechanical perturbation support the pit area or conduit reinforcement hypotheses in Phaseolus vulgaris?

Two Phaseolus vulgaris L. cultivars were exposed to reduced water and stem mechanical perturbation treatments (flexing) to determine if acclimation to these treatments induced hydraulic changes, altered cavitation resistance and changed stem mechanical properties. Additionally, this study sought to determine if changes in cavitation resistance would support the pit area or conduit reinforcement hypotheses. Flexing reduced biomass, leaf area, xylem vessel area and hydraulic conductivity. One cultivar had greater measures of stem strength and cavitation resistance. Flexing increased cavitation resistance (P50 ) but did not increase Young's modulus, rigidity or flexural strength on dried stems. Stem rigidity and basal diameter were correlated with leaf mass. The ratio of conduit wall thickness to span [(t/b)h 2 ] increased under high water and flexing treatments while rigidity decreased for one cultivar exposed to both flexing and lower water suggesting an inability to compensate for two simultaneous stresses. Although P50 was not correlated with measures of mechanical strength, P50 was correlated with vessel diameter, consistent with the pit area hypothesis. This study confirmed that mechanical perturbation can impact xylem structural properties and result in altered plant water flow characteristics and cavitation resistance. Long-term hydraulic acclimation in these herbaceous annuals was constrained by similar tradeoffs that constrain hydraulic properties across species.

Constraints to hydraulic acclimation under reduced light in two contrasting Phaseolus vulgaris cultivars.

Two cultivars of Phaseolus vulgaris L. were grown under three light levels to determine if hydraulic acclimation to light occurs in herbaceous annuals and whether intraspecific trade-offs constrain hydraulic traits. Acclimation occurred in response to reduced light and included decreased stomatal density (SD) and increased specific leaf area (SLA). Reduced light resulted in lower wood density (WD); decreased cavitation resistance, measured as the xylem pressure causing a 50 % reduction in stem conductivity (P50); and increased hydraulic capacity, measured as average leaf mass specific transpiration (E(LM)). Significant or marginally significant trade-offs between P50 and WD, WD and E(LM), and E(LM) and P50 reflected variation due to both genotype and environmental effects. A trade-off between WD and P50 within one cultivar indicated that morphological adjustment was constrained. Coordinated changes in WD, P50, and E(LM) within each cultivar in response to light were consistent with trade-offs constraining plasticity. A water-use efficiency (WUE, measured as δ(13)C) versus hydraulic capacity (E(LM)) trade-off was observed within each cultivar, further indicating that hydraulic trade-offs can constrain acclimation. Larger plants had lower hydraulic capacity (E(LM)) but greater cavitation resistance, WD, and WUE. Distinct hydraulic strategies were observed with the cultivar adapted to irrigated conditions having higher stomatal conductance and stem flow rates. The cultivar adapted to rain-fed conditions had higher leaf area and greater cavitation resistance. Hydraulic trade-offs were observed within the herbaceous P. vulgaris resulting from both genotype and environmental effects. Trade-offs within a cultivar reflected constraints to hydraulic acclimation in response to changing light.

Evolutionary constraint on low elevation range expansion: Defense-abiotic stress-tolerance trade-off in crosses of the ecological model Boechera stricta.

Most transplant experiments across species geographic range boundaries indicate that adaptation to stressful environments outside the range is often constrained. However, the mechanisms of these constraints remain poorly understood. We used extended generation crosses from diverged high and low elevation populations. In experiments across low elevation range boundaries, there was selection on the parental lines for abiotic stress-tolerance and resistance to herbivores. However, in support of a defense-tolerance trade-off, extended generation crosses showed nonindependent segregation of these traits in the laboratory across a drought-stress gradient and in the field across the low elevation range boundary. Genotypic variation in a marker from a region of the genome containing a candidate gene (MYC2) was associated with change in the genetic trade-off. Thus, using crosses and forward genetics, we found experimental genetic and molecular evidence for a pleiotropic trade-off that could constrain the evolution of range expansion.

Effect of Drought on Herbivore-Induced Plant Gene Expression: Population Comparison for Range Limit Inferences.

Low elevation "trailing edge" range margin populations typically face increases in both abiotic and biotic stressors that may contribute to range limit development. We hypothesize that selection may act on ABA and JA signaling pathways for more stable expression needed for range expansion, but that antagonistic crosstalk prevents their simultaneous co-option. To test this hypothesis, we compared high and low elevation populations of Boechera stricta that have diverged with respect to constitutive levels of glucosinolate defenses and root:shoot ratios; neither population has high levels of both traits. If constraints imposed by antagonistic signaling underlie this divergence, one would predict that high constitutive levels of traits would coincide with lower plasticity. To test this prediction, we compared the genetically diverged populations in a double challenge drought-herbivory growth chamber experiment. Although a glucosinolate defense response to the generalist insect herbivore Spodoptera exigua was attenuated under drought conditions, the plastic defense response did not differ significantly between populations. Similarly, although several potential drought tolerance traits were measured, only stomatal aperture behavior, as measured by carbon isotope ratios, was less plastic as predicted in the high elevation population. However, RNAseq results on a small subset of plants indicated differential expression of relevant genes between populations as predicted. We suggest that the ambiguity in our results stems from a weaker link between the pathways and the functional traits compared to transcripts.

Patterns of stomatal conductance among blue oak (Quercus douglasii) size classes and populations: implications for seedling establishment.

Quercus douglasii Hook. & Arn. (blue oak) is a deciduous white oak that is currently failing to regenerate throughout much of its range in California, USA. Patterns of water use were observed in adult trees, saplings and seedlings to determine if ontogenetic changes in water use occur, which might be important in the establishment of this long-lived perennial species in a Mediterranean-type system. Seasonal and diurnal stomatal conductance (g(s)), late-season predawn xylem water potentials (Psi(pre)), carbon isotopic ratio (delta(13)C) and soil water status were compared among the three size classes at three sites differing in mean precipitation and soil water characteristics. Comparisons were also made between microsites with and without regeneration (defined by the presence or absence of saplings). Overall patterns of water use were consistent among the three sites, except that, at the site with the highest rainfall, Q. douglasii plants had higher g(s) and more positive Psi(pre) values. Although no differences in water use patterns were found between regeneration and non-regeneration microsites, the observed ontogenetic differences in water use may have important implications for Q. douglasii establishment. Compared with adult trees and saplings, seedlings had higher gas exchange rates during periods of high soil water content (early in the season and in the morning). Seedling g(s) was correlated with percent extractable soil water (ESW) throughout the season; adult tree and sapling g(s) was correlated with ESW between June and September. Despite experiencing greater water stress (indicated by more negative Psi(pre) values) than older trees, seedlings had more negative delta(13)C values, implying lower water-use efficiencies.

Drought-induced trans-generational tradeoff between stress tolerance and defence: consequences for range limits?

Areas just across species range boundaries are often stressful, but even with ample genetic variation within and among range-margin populations, adaptation towards stress tolerance across range boundaries often does not occur. Adaptive trans-generational plasticity should allow organisms to circumvent these problems for temporary range expansion; however, range boundaries often persist. To investigate this dilemma, we drought stressed a parent generation of Boechera stricta (A.Gray) A. Löve & D. Löve, a perennial wild relative of Arabidopsis, representing genetic variation within and among several low-elevation range margin populations. Boechera stricta is restricted to higher, moister elevations in temperate regions where generalist herbivores are often less common. Previous reports indicate a negative genetic correlation (genetic tradeoff) between chemical defence allocation and abiotic stress tolerance that may prevent the simultaneous evolution of defence and drought tolerance that would be needed for range expansion. In growth chamber experiments, the genetic tradeoff became undetectable among offspring sib-families whose parents had been drought treated, suggesting that the stress-induced trans-generational plasticity may circumvent the genetic tradeoff and thus enable range expansion. However, the trans-generational effects also included a conflict between plastic responses (environmental tradeoff); offspring whose parents were drought treated were more drought tolerant, but had lower levels of glucosinolate toxins that function in defence against generalist herbivores. We suggest that either the genetic or environmental tradeoff between defence allocation and stress tolerance has the potential to contribute to range limit development in upland mustards.

Long-term acclimatization of hydraulic properties, xylem conduit size, wall strength and cavitation resistance in Phaseolus vulgaris in response to different environmental effects.

Phaseolus vulgaris grown under various environmental conditions was used to assess long-term acclimatization of xylem structural characteristics and hydraulic properties. Conduit diameter tended to be reduced and 'wood' density (of 'woody' stems) increased under low moisture ('dry'), increased soil porosity ('porous soil') and low phosphorus ('low P') treatments. Dry and low P had the largest percentage of small vessels. Dry, low light ('shade') and porous soil treatments decreased P50 (50% loss in conductivity) by 0.15-0.25 MPa (greater cavitation resistance) compared with 'controls'. By contrast, low P increased P50 by 0.30 MPa (less cavitation resistance) compared with porous soil (the control for low P). Changes in cavitation resistance were independent of conduit diameter. By contrast, changes in cavitation resistance were correlated with wood density for the control, dry and porous soil treatments, but did not appear to be a function of wood density for the shade and low P treatments. In a separate experiment comparing control and porous soil plants, stem hydraulic conductivity (kh), specific conductivity (ks), leaf specific conductivity (LSC), total pot water loss, plant biomass and leaf area were all greater for control plants compared to porous soil plants. Porous soil plants, however, demonstrated higher midday stomatal conductance to water vapour (gs), apparently because they experienced proportionally less midday xylem cavitation.

Patterns of tree dieback in Queensland, Australia: the importance of drought stress and the role of resistance to cavitation.

During the extreme 1992-1997 El Niño drought event, widespread stem mortality, or tree "dieback", of both mature and juvenile eucalypts occurred within the tropical savannas of northeast Australia. Most of the dieback occurred in individuals of the ironbark species complex ( Eucalyptus crebra- E. xanthoclada) while individuals of the bloodwood species Corymbia erythrophloia, exhibited significantly less stem mortality. Indicative of greater water stress, predawn and midday xylem water potentials of ironbark adults and saplings were significantly more negative than predawn values of bloodwoods. The very negative xylem water potentials in ironbarks suggest that stem mortality in both adult and juvenile ironbarks results from drought-induced embolism and that ironbarks perhaps have a shallower and less extensive root system than bloodwoods. Although predawn and midday water potentials for ironbark adults and saplings were similar, a census of mature and juvenile ironbark trees indicated that mortality was higher in adult trees. Cavitation vulnerability curves indicated that ironbark saplings may be better buffered against cavitation than adult trees. If they possess smaller root systems, saplings are more likely than adults to experience low xylem water potentials, even in non-drought years. Xylem conduits produced in adult trees during periods of normal rainfall, although perhaps more efficient in water conduction, may be more vulnerable to cavitation during infrequent severe droughts.