Low root reserve accumulation during drought may lead to winter mortality in poplar seedlings.

Climate models suggest that more frequent drought events of greater severity and length, associated with climate change, can be expected in the coming decades. Although drought-induced tree mortality has been recognized as an important factor modulating forest demography at the global scale, the mechanisms underlying drought-induced tree mortality remain contentious. Above- and below-ground growth, gas exchange, water relations and carbon reserve accumulation dynamics at the organ and whole-plant scale were quantified in Populus tremuloides and P. balsamifera seedlings in response to severe drought. Seedlings were maintained in drought conditions over one growing and one dormant winter season. Our experiment presents a detailed description of the effect of severe drought on growth and physiological variables, leading to seedling mortality after an extended period of drought and dormancy. After re-watering following the dormant period, drought-exposed seedlings did not re-flush, showing that the root system had died off. The results of this study suggest a complex series of physiological feedbacks between the measured variables in both Populus species. Further, they reveal that reduced reserve accumulation in the root system during drought decreases the conversion of starch to soluble sugars in roots, which may contribute to the root death of drought-exposed seedlings during the dormant season by compromising the frost tolerance of the root system.

Impact of simulated herbivory on water relations of aspen (Populus tremuloides) seedlings: the role of new tissue in the hydraulic conductivity recovery cycle.

Physiological mechanisms behind plant-herbivore interactions are commonly approached as input-output systems where the role of plant physiology is viewed as a black box. Studies evaluating impacts of defoliation on plant physiology have mostly focused on changes in photosynthesis while the overall impact on plant water relations is largely unknown. Stem hydraulic conductivity (k(h)), stem specific conductivity (k(s)), percent loss of hydraulic conductivity (PLC), CO(2) assimilation (A) and stomatal conductance (g(s)) were measured on well-irrigated 1-month-old Populus tremuloides (Michx.) defoliated and control seedlings until complete refoliation. PLC values of defoliated seedlings gradually increased during the refoliation process despite them being kept well irrigated. k(s) of defoliated seedlings gradually decreased during refoliation. PLC and k(s) values of control seedlings remained constant during refoliation. k(s) of new stems, leaf specific conductivity and A of leaves grown from new stems in defoliated and control seedlings were not significantly different, but g(s) was higher in defoliated than in control seedlings. The gradual increase of PLC and decrease of k(s) values in old stems after defoliation was unexpected under well-irrigated conditions, but appeared to have little impact on new stems formed after defoliation. The gradual loss of conductivity measured during the refoliation process under well-irrigated conditions suggests that young seedlings of P. tremuloides may be more susceptible to cavitation after herbivore damage under drought conditions.

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Resource availability and repeated defoliation mediate compensatory growth in trembling aspen (Populus tremuloides) seedlings.

Plant ecologists have debated the mechanisms used by plants to cope with the impact of herbivore damage. While plant resistance mechanisms have received much attention, plant compensatory growth as a type of plant tolerance mechanisms has been less studied. We conducted a greenhouse experiment to evaluate compensatory growth for trembling aspen (Populus tremuloides) seedlings under varying intensities and frequencies of simulated defoliation, with or without nutrient enriched media. For the purpose of this study, changes in biomass production and non-structural carbohydrate concentrations (NSC) of roots and leaves were considered compensatory responses. All defoliated seedlings showed biomass accumulation under low defoliation intensity and frequency, regardless of resource availability; however, as defoliation intensity and frequency increased, compensatory growth of seedlings was altered depending on resource availability. Seedlings in a resource-rich environment showed complete compensation, in contrast responses ranged from undercompensation to complete compensation in a resource-limited environment. Furthermore, at the highest defoliation intensity and frequency, NSC concentrations in leaves and roots were similar between defoliated and non-defoliated seedlings in a resource-rich environment; in contrast, defoliated seedlings with limited resources sustained the most biomass loss, had lower amounts of stored NSC. Using these results, we developed a new predictive framework incorporating the interactions between frequency and intensity of defoliation and resource availability as modulators of plant compensatory responses.

Root carbon reserve dynamics in aspen seedlings: does simulated drought induce reserve limitation?

In a greenhouse study we quantified the gradual change of gas exchange, water relations and root reserves of aspen (Populus tremuloides Michx.) seedlings growing over a 3-month period of severe water stress. The aim of the study was to quantify the complex interrelationship between growth, water and gas exchange, and root carbon (C) dynamics. Various growth, gas exchange and water relations variables in combination with root reserves were measured periodically on seedlings that had been exposed to a continuous drought treatment over a 12-week period and compared with well-watered seedlings. Although gas exchange and water relations parameters significantly decreased over the drought period in aspen seedlings, root reserves did not mirror this trend. During the course of the experiment roots of aspen seedlings growing under severe water stress showed a two orders of magnitude increase in sugar and starch content, and roots of these seedlings contained more starch relative to sugar than those in non-droughted seedlings. Drought resulted in a switch from growth to root reserves storage which indicates a close interrelationship between growth and physiological variables and the accumulation of root carbohydrate reserves. Although a severe 3-month drought period created physiological symptoms of C limitation, there was no indication of a depletion of root C reserve in aspen seedlings.

Short dry spells in the wet season increase mortality of tropical pioneer seedlings.

Variation in plant species performance in response to water availability offers a potential axis for temporal and spatial habitat partitioning and may therefore affect community composition in tropical forests. We hypothesized that short dry spells during the wet season are a significant source of mortality for the newly emerging seedlings of pioneer species that recruit in treefall gaps in tropical forests. An analysis of a 49-year rainfall record for three forests across a rainfall gradient in central Panama confirmed that dry spells of > or = 10 days during the wet season occur on average once a year in a deciduous forest, and once every other year in a semi-deciduous moist and an evergreen wet forest. The effect of wet season dry spells on the recruitment of pioneers was investigated by comparing seedling survival in rain-protected dry plots and irrigated control plots in four large artificially created treefall gaps in a semi-deciduous tropical forest. In rain-protected plots surface soil layers dried rapidly, leading to a strong gradient in water potential within the upper 10 cm of soil. Seedling survival for six pioneer species was significantly lower in rain-protected than in irrigated control plots after only 4 days. The strength of the irrigation effect differed among species, and first became apparent 3-10 days after treatments started. Root allocation patterns were significantly, or marginally significantly, different between species and between two groups of larger and smaller seeded species. However, they were not correlated with seedling drought sensitivity, suggesting allocation is not a key trait for drought sensitivity in pioneer seedlings. Our data provide strong evidence that short dry spells in the wet season differentially affect seedling survivorship of pioneer species, and may therefore have important implications to seedling demography and community dynamics.