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• 1. [Article] Physiological Responses of Loblolly Pine and Douglas-fir Seedlings from Various Provenances to Timing and Frequency of Drought Stress

  Drought is expected to increase in many parts of the world and has been shown to affect tree physiology and growth, with seedlings being particularly vulnerable. Seedling drought responses are often species ...

  Citation × Citation Citation Abstract Copy Title: Physiological Responses of Loblolly Pine and Douglas-fir Seedlings from Various Provenances to Timing and Frequency of Drought Stress Author: Mosel, Jamie E. Drought is expected to increase in many parts of the world and has been shown to affect tree physiology and growth, with seedlings being particularly vulnerable. Seedling drought responses are often species dependent, and even within species different populations may demonstrate a spectrum of responses to drought, from susceptibility to resistance. As both loblolly pine (Pinus taeda) and Douglas-fir (Pseudotsuga menziesii) have broad geographic ranges, they provide an opportunity to assess drought resistances of provenances associated with a variety of site climate conditions across their ranges. Furthermore, while studies of singular drought events have revealed important information about seedling stress responses, it has been shown that drought may impart effects even following the release from drought. In some cases, effects of drought exposure can be detrimental or lethal; however, exposure to non-lethal drought may also lead to drought acclimation, which could potentially enhance seedling functioning under subsequent droughts. This potential phenotypic plasticity, i.e. the extent to which a seedling can acclimate to drought conditions, may vary across species and across provenances within a species. This study used physiological (electron transport rate, fluorescence, and water potentials) and growth (biomass accumulation and height increment) responses for two goals: 1) to assess the drought resistances of three provenances of loblolly pine and three provenances of Douglas-fir associated with varying site climate conditions, and 2) to investigate whether previous exposure to a drought results in acclimation to a second drought. We hypothesized that: 1) drought responses would differ among the provenances in accordance with associated site climate conditions (i.e., provenances associated with mesic site climates as characterized by low climatic moisture deficit and high mean annual precipitation would be drought susceptible, and more xeric site climates as characterized by high climatic moisture deficit and low mean annual precipitation would be more drought resistant) as evidenced by physiological measures of electron transport rate, fluorescence, water potentials and growth; and 2) previous exposure to drought would result in acclimation to drought as evidenced by maintenance of physiological function (i.e., higher levels of electron transport and fluorescence) in previously drought exposed seedlings compared to previously unexposed seedlings. We also hypothesized that levels of acclimation would vary among provenances. The study yielded some evidence to support the first hypothesis regarding provenance differences in drought resistances in both species. Provenance drought resistances conformed largely to expectations, though differences were less than expected. In loblolly pine, although not statistically significant, during the second drought there was a pattern of lower maximum electron transport rates, which appeared sooner in the more mesic provenance than in the other two provenances. There were also provenance differences in seedling heights, with the most xeric of the three provenances being shorter at the beginning and the end of the study. In Douglas-fir, there were significant differences in provenance and in the interaction of treatment and provenance for maximum electron rates and fluorescence. Dark-adapted fluorescence was lower in the Coos Bay (mesic site climate) provenance during drought than in the Cascades (mesic site climate) and New Mexico (xeric site climate) provenances. The New Mexico provenance showed the least differences in fluorescence between droughted and watered treatments. The study also yielded some evidence to support the second hypothesis regarding drought acclimation in both species and among provenances. During the second drought, there was a pattern of higher maximum electron transport rates and fluorescence in previously drought exposed seedlings as compared to seedlings previously unexposed to drought. This difference was significant in the mesic provenance (North Carolina) of loblolly pine, with maximum electron transport rates significantly higher in the previously drought exposed treatment compared to the newly exposed treatment during and following the second drought. Patterns of lower electron transport rates in seedlings previously unexposed to drought compared to seedlings previously exposed to drought also appeared in Douglas-fir during the second drought, though not with statistical significance. However, the most xeric provenance showed the reverse pattern during and following the second drought, with lower maximum electron transport rates in the previously drought exposed treatment compared to the treatment previously unexposed to drought. Chlorophyll fluorescence values were significantly higher during the second drought in the previously droughted treatments compared to newly drought-exposed treatments in some provenances of loblolly pine and Douglas-fir. Lastly, seedlings exposed to an early drought had significantly lower final heights than seedlings unexposed to an early drought in both species, although the differences were greater in loblolly pine than Douglas-fir. It may be that growth acclimation, especially in the form of height reductions, influenced physiological responses during a second drought. Further studies are necessary to provide more conclusive evidence in support or against the two hypotheses. Nonetheless, this study provides valuable information on the drought responses of young, greenhouse-grown seedlings of two species that are widespread in North America and that are economically important throughout the world. Further studies in a wider range of age classes, incorporating field studies or more natural settings, may help better predict plant responses in the face of changing climate. Title: Physiological Responses of Loblolly Pine and Douglas-fir Seedlings from Various Provenances to Timing and Frequency of Drought Stress Author: Mosel, Jamie E. Drought is expected to increase in many parts of the world and has been shown to affect tree physiology and growth, with seedlings being particularly vulnerable. Seedling drought responses are often species dependent, and even within species different populations may demonstrate a spectrum of responses to drought, from susceptibility to resistance. As both loblolly pine (Pinus taeda) and Douglas-fir (Pseudotsuga menziesii) have broad geographic ranges, they provide an opportunity to assess drought resistances of provenances associated with a variety of site climate conditions across their ranges. Furthermore, while studies of singular drought events have revealed important information about seedling stress responses, it has been shown that drought may impart effects even following the release from drought. In some cases, effects of drought exposure can be detrimental or lethal; however, exposure to non-lethal drought may also lead to drought acclimation, which could potentially enhance seedling functioning under subsequent droughts. This potential phenotypic plasticity, i.e. the extent to which a seedling can acclimate to drought conditions, may vary across species and across provenances within a species. This study used physiological (electron transport rate, fluorescence, and water potentials) and growth (biomass accumulation and height increment) responses for two goals: 1) to assess the drought resistances of three provenances of loblolly pine and three provenances of Douglas-fir associated with varying site climate conditions, and 2) to investigate whether previous exposure to a drought results in acclimation to a second drought. We hypothesized that: 1) drought responses would differ among the provenances in accordance with associated site climate conditions (i.e., provenances associated with mesic site climates as characterized by low climatic moisture deficit and high mean annual precipitation would be drought susceptible, and more xeric site climates as characterized by high climatic moisture deficit and low mean annual precipitation would be more drought resistant) as evidenced by physiological measures of electron transport rate, fluorescence, water potentials and growth; and 2) previous exposure to drought would result in acclimation to drought as evidenced by maintenance of physiological function (i.e., higher levels of electron transport and fluorescence) in previously drought exposed seedlings compared to previously unexposed seedlings. We also hypothesized that levels of acclimation would vary among provenances. The study yielded some evidence to support the first hypothesis regarding provenance differences in drought resistances in both species. Provenance drought resistances conformed largely to expectations, though differences were less than expected. In loblolly pine, although not statistically significant, during the second drought there was a pattern of lower maximum electron transport rates, which appeared sooner in the more mesic provenance than in the other two provenances. There were also provenance differences in seedling heights, with the most xeric of the three provenances being shorter at the beginning and the end of the study. In Douglas-fir, there were significant differences in provenance and in the interaction of treatment and provenance for maximum electron rates and fluorescence. Dark-adapted fluorescence was lower in the Coos Bay (mesic site climate) provenance during drought than in the Cascades (mesic site climate) and New Mexico (xeric site climate) provenances. The New Mexico provenance showed the least differences in fluorescence between droughted and watered treatments. The study also yielded some evidence to support the second hypothesis regarding drought acclimation in both species and among provenances. During the second drought, there was a pattern of higher maximum electron transport rates and fluorescence in previously drought exposed seedlings as compared to seedlings previously unexposed to drought. This difference was significant in the mesic provenance (North Carolina) of loblolly pine, with maximum electron transport rates significantly higher in the previously drought exposed treatment compared to the newly exposed treatment during and following the second drought. Patterns of lower electron transport rates in seedlings previously unexposed to drought compared to seedlings previously exposed to drought also appeared in Douglas-fir during the second drought, though not with statistical significance. However, the most xeric provenance showed the reverse pattern during and following the second drought, with lower maximum electron transport rates in the previously drought exposed treatment compared to the treatment previously unexposed to drought. Chlorophyll fluorescence values were significantly higher during the second drought in the previously droughted treatments compared to newly drought-exposed treatments in some provenances of loblolly pine and Douglas-fir. Lastly, seedlings exposed to an early drought had significantly lower final heights than seedlings unexposed to an early drought in both species, although the differences were greater in loblolly pine than Douglas-fir. It may be that growth acclimation, especially in the form of height reductions, influenced physiological responses during a second drought. Further studies are necessary to provide more conclusive evidence in support or against the two hypotheses. Nonetheless, this study provides valuable information on the drought responses of young, greenhouse-grown seedlings of two species that are widespread in North America and that are economically important throughout the world. Further studies in a wider range of age classes, incorporating field studies or more natural settings, may help better predict plant responses in the face of changing climate. Close