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Abstract A two decade‐long megadrought, with likely anthropogenic causes, has impacted forest growth and mortality across the southwestern U.S. Given this event, and the future likelihood of similar climate challenges, it is important to understand how different water resources are used by semi‐arid forests in this region. Within the geographic domain of the North American Monsoon climate system, we studied seasonal water‐use in eight differentPinus ponderosamontane forests distributed across a climate gradient with varying contributions from winter and summer precipitation. We collected oxygen isotopes from precipitation, soil, and xylem water during two contrasting hydrologic years to determine how trees differentially use winter versus summer precipitation sources. Most trees switched from using snowmelt water as the primary source during the early‐summer hyper‐arid period, to monsoon rainwater during the late‐summer. However, during the low snowpack year, which represents the most common climate phenomenon during the megadrought, trees at all sites used less summer rain when compared to the higher snowpack year, demonstrating a drought‐induced antecedent influence of winter precipitation on the uptake of summer rain. A possible mechanism to explain the antecedent effect is an earlier snow disappearance during the low snowpack year weakening hydrologic connectivity within the soil profile, decreasing the soil infiltration of summer rains. However, in years with higher snowpack, the snow lasts longer, and this can improve the hydrologic connectivity within the soil profile. As a result, there is more infiltration of summer rains into the soils. This can enhance the maintenance of active shallow fine‐root biomass during the period when snowpack disappears, and monsoon rains have yet to arrive. These findings provide insight into how the seasonal interactions between major seasonal climate systems influence forest tree water use in the face of an extreme megadrought. 

Abstract The US Southwest has been entrenched in a two‐decade‐long megadrought (MD), the most severe since 800 CE, which threatens the long‐term vitality and persistence of regional montane forests. Here, we report that in the face of record low winter precipitation and increasing atmospheric aridity, seasonal activity of the North American Monsoon (NAM) climate system brings sufficient precipitation during the height of the summer to alleviate extreme tree water stress. We studied seasonally resolved, tree‐ring stable carbon isotope ratios across a 57‐year time series (1960–2017) in 17 Ponderosa pine forests distributed across the NAM geographic domain. Our study focused on the isotope dynamics of latewood (LW), which is produced in association with NAM rains. During the MD, populations growing within the core region of the NAM operated at lower intrinsic and higher evaporative water‐use efficiencies (WUE_iand WUE_E, respectively), compared to populations growing in the periphery of the NAM domain, indicating less physiological water stress in those populations with access to NAM moisture. The disparities in water‐use efficiencies in periphery populations are due to a higher atmospheric vapor pressure deficit (VPD) and reduced access to summer soil moisture. The buffering advantage of the NAM, however, is weakening. We observed that since the MD, the relationship between WUE_iand WUE_Ein forests within the core NAM domain is shifting toward a drought response similar to forests on the periphery of the NAM. After correcting for past increases in the atmospheric CO₂concentration, we were able to isolate the LW time‐series responses to climate alone. This showed that the shift in the relation between WUE_iand WUE_Ewas driven by the extreme increases in MD‐associated VPD, with little advantageous influence on stomatal conductance from increases in atmospheric CO₂concentration. 

Abstract Tree‐ring carbon and oxygen isotope ratios have been used to understand past dynamics in forest carbon and water cycling. Recently, this has been possible for different parts of single growing seasons by isolating anatomical sections within individual annual rings. Uncertainties in this approach are associated with correlated climate legacies that can occur at a higher frequency, such as across successive seasons, or a lower frequency, such as across years. The objective of this study was to gain insight into how legacies affect cross‐correlation in the δ¹³C and δ¹⁸O isotope ratios in the earlywood (EW) and latewood (LW) fractions ofPinus ponderosatrees at thirteen sites across a latitudinal gradient influenced by the North American Monsoon (NAM) climate system. We observed that δ¹³C from EW and LW has significant positive cross‐correlations at most sites, whereas EW and LW δ¹⁸O values were cross‐correlated at about half the sites. Using combined statistical and mechanistic models, we show that cross‐correlations in both δ¹³C and δ¹⁸O can be largely explained by a low‐frequency (multiple‐year) mode that may be associated with long‐term climate change. We isolated, and statistically removed, the low‐frequency correlation, which resulted in greater geographical differentiation of the EW and LW isotope signals. The remaining higher‐frequency (seasonal) cross‐correlations between EW and LW isotope ratios were explored using a mechanistic isotope fractionation–climate model. This showed that lower atmospheric vapor pressure deficits associated with monsoon rain increase the EW‐LW differentiation for both δ¹³C and δ¹⁸O at southern sites, compared to northern sites. Our results support the hypothesis that dominantly unimodal precipitation regimes, such as near the northern boundary of the NAM, are more likely to foster cross‐correlations in the isotope signals of EW and LW, potentially due to greater sharing of common carbohydrate and soil water resource pools, compared to southerly sites with bimodal precipitation regimes. 

Recent evidence has revealed the emergence of a megadrought in southwestern North America since 2000. Megadroughts extend for at least 2 decades, making it challenging to identify such events until they are well established. Here, we examined tree-ring growth and stable isotope ratios in Pinus ponderosa at its driest niche edge to investigate whether trees growing near their aridity limit were sensitive to the megadrought climatic pre-conditions, and were capable of informing predictive efforts. During the decade before the megadrought, trees in four populations revealed increases in the cellulose δ13C content of earlywood, latewood, and false latewood, which, based on past studies are correlated with increased intrinsic water-use efficiency. However, radial growth and cellulose δ18O were not sensitive to pre-megadrought conditions. During the 2 decades preceding the megadrought, at all four sites, the changes in δ13C were caused by the high sensitivity of needle carbon and water exchange to drought trends in key winter months, and for three of the four sites during crucial summer months. Such pre-megadrought physiological sensitivity appears to be unique for trees near their arid range limit, as similar patterns were not observed in trees in ten reference sites located along a latitudinal gradient in the same megadrought domain, despite similar drying trends. Our results reveal the utility of tree-ring δ13C to reconstruct spatiotemporal patterns during the organizational phase of a megadrought, demonstrating that trees near the arid boundaries of a species’ distribution might be useful in the early detection of long-lasting droughts. 

Plant isoprene emissions are known to contribute to abiotic stress tolerance, especially during episodes of high temperature and drought, and during cellular oxidative stress. Recent studies have shown that genetic transformations to add or remove isoprene emissions cause a cascade of cellular modifications that include known signaling pathways, and interact to remodel adaptive growth-defense tradeoffs. The most compelling evidence for isoprene signaling is found in the shikimate and phenylpropanoid pathways, which produce salicylic acid, alkaloids, tannins, anthocyanins, flavonols and other flavonoids; all of which have roles in stress tolerance and plant defense. Isoprene also influences key gene expression patterns in the terpenoid biosynthetic pathways, and the jasmonic acid, gibberellic acid and cytokinin signaling networks that have important roles in controlling inducible defense responses and influencing plant growth and development, particularly following defoliation. In this synthesis paper, using past studies of transgenic poplar, tobacco and Arabidopsis, we present the evidence for isoprene acting as a metabolite that coordinates aspects of cellular signaling, resulting in enhanced chemical defense during periods of climate stress, while minimizing costs to growth. This perspective represents a major shift in our thinking away from direct effects of isoprene, for example, by changing membrane properties or quenching ROS, to indirect effects, through changes in gene expression and protein abundances. Recognition of isoprene’s role in the growth-defense tradeoff provides new perspectives on evolution of the trait, its contribution to plant adaptation and resilience, and the ecological niches in which it is most effective. 

Hybrid-poplar tree plantations provide a source for biofuel and biomass, but they also increase forest isoprene emissions. The consequences of increased isoprene emissions include higher rates of tropospheric ozone production, increases in the lifetime of methane, and increases in atmospheric aerosol production, all of which affect the global energy budget and/or lead to the degradation of air quality. Using RNA interference (RNAi) to suppress isoprene emission, we show that this trait, which is thought to be required for the tolerance of abiotic stress, is not required for high rates of photosynthesis and woody biomass production in the agroforest plantation environment, even in areas with high levels of climatic stress. Biomass production over 4 y in plantations in Arizona and Oregon was similar among genetic lines that emitted or did not emit significant amounts of isoprene. Lines that had substantially reduced isoprene emission rates also showed decreases in flavonol pigments, which reduce oxidative damage during extremes of abiotic stress, a pattern that would be expected to amplify metabolic dysfunction in the absence of isoprene production in stress-prone climate regimes. However, compensatory increases in the expression of other proteomic components, especially those associated with the production of protective compounds, such as carotenoids and terpenoids, and the fact that most biomass is produced prior to the hottest and driest part of the growing season explain the observed pattern of high biomass production with low isoprene emission. Our results show that it is possible to reduce the deleterious influences of isoprene on the atmosphere, while sustaining woody biomass production in temperate agroforest plantations.