Lisa A. Donovan

Nighttime Stomatal Conductance and Transpiration in C3 and C4 Plants

Incomplete stomatal closure during the night is observed in a diverse range of C3 and C4 species ([Fig. 1][1] ; Supplemental Table S1) and can lead to substantial nighttime transpirational water loss. Although water loss is an inevitable consequence of stomatal opening for photosynthetic carbon gain

Water potential and ionic effects on germination and seedling growth of two cold desert shrubs

We tested expectations that two desert shrubs would differ in germination and seedling relative growth rate (RGR) responses to Na and Ψ(s) stress. The study species, Chrysothamnus nauseosus ssp. consimilis and Sarcobatus vermiculatus (hereafter referred to by genus), differ in their distribution along salinity gradients, with Chrysothamnus inhabiting only less saline areas. In growth chamber studies, declining Ψ(s) (-0.82 to -2.71 MPa) inhibited germination of both species, and Chrysothamnus was less tolerant of Ψ(s) stress than Sarcobatus. Germination fell below 10% for Chrysothamnus at -1.64 MPa (NaCl and PEG), and for Sarcobatus at -2.4 MPa PEG. Neither species exhibited ion toxicity. There was substantial ion enhancement for Sarcobatus in lower Ψ(s), allowing for 40% germination in -2.71 MPa NaCl. For seedling RGR, species were not different at -0.29 or -0.82 MPa (0 and 100 mmol/L NaCl, respectively), but Chrysothamnus RGR declined substantially at -1.3 MPa (200 mmol/L NaCl). The greater stress tolerance of Sarcobatus was not associated with a lower RGR under nonsaline conditions. Species differences in seed and seedling Ψ(s) stress tolerance probably contribute to the restricted distribution of Chrysothamnus to less saline areas. The Na uptake of Sarcobatus seedlings enhances its ability to deal with declining Ψ(s) and establish in more saline areas.

Magnitude and mechanisms of disequilibrium between predawn plant and soil water potentials

Predawn plant water potential (Ψw, measured with leaf psychrometers) and surrogate measurements made with the pressure chamber (termed Ψpc here) are used to infer comparative ecological performance, based on the expectation that these plant potentials reflect the wettest soil Ψw accessed by roots. There is growing evidence, however, that some species exhibit substantial predawn disequilibrium (PDD), defined as plant Ψw or Ψpc at predawn substantially more negative than the Ψw of soil accessed by roots. In the western Great Basin desert, the magnitude of PDD calculated as soil Ψw minus predawn leaf Ψw was as large as 1.4 and 2.7 MPa for two codominant shrub species, Chrysothamnus nauseosus and Sarcobatus vermiculatus, respectively. The magnitude of PDD calculated as soil Ψw minus predawn Ψpc was smaller, up to 0.6 and 2.1 MPa for Chrysothamnus and Sarcobatus, respectively. For both species, mechanisms contributing to PDD included nighttime transpiration and putative leaf apoplastic solutes, but not hydraul...

Transgressive character expression in a hybrid sunflower species

Diploid hybrid lineages often are ecologically distinct from their parental species. However, it is unclear whether this niche divergence is typically achieved via hybrid intermediacy, a mixture of parental traits, and/or the evolution of extreme (transgressive) morphological and ecophysiological features. Here we compare an extensively studied hybrid sunflower species, Helianthus anomalus, with its putative parents, H. annuus and H. petiolaris, for 41 morphological and 12 ecophysiological traits. Helianthus anomalus was morphologically intermediate for one trait (2.4%), parental-like for 23 traits (56.1%), and transgressive for 17 traits (41.5%). For ecophysiological traits, H. anomalus was not significantly different from one or both parents for nine traits (75%), and was transgressive for the remaining three (25%). Thus, H. anomalus appears to be a mosaic of parental-like and transgressive phenotypes. Although the fitness effects of the transgressive characters are not yet known, many of these characters are consistent with adaptations reported for other sand dune plants. Genetic studies are currently underway to ascertain whether these extreme characters arose as a direct byproduct of hybridization or whether they evolved via mutational divergence.

The evolution of the worldwide leaf economics spectrum

The worldwide leaf economic spectrum (WLES) is a strikingly consistent pattern of correlations among leaf traits. Although the WLES effectively summarizes variation in plant ecological strategies, little is known about its evolution. We reviewed estimates of natural selection and genetic variation for leaf traits to test whether the evolution of the WLES was limited by selection against unfit trait combinations or by genetic constraints. There was significant selection for leaf traits on both ends of the WLES spectrum, as well as significant genetic variation for these traits. In addition, genetic correlations between WLES traits were variable in strength and direction. These data suggest that genetic constraints have had a smaller role than selection in the evolution of the WLES.

Genetic variation in Pueraria lobata (Fabaceae), an introduced, clonal, invasive plant of the southeastern United States

Pueraria lobata (kudzu), a clonal, leguminous vine, is invading the southeastern United States at a rate of 50 000 ha per year. Genetic variability and clonal diversity were measured in 20 southeastern U.S. populations using 14 allozyme loci. Within its U.S. range, 92.9% of the loci were polymorphic and overall genetic diversity was 0.290. Such high levels of genetic diversity are consistent with its history of multiple introductions over an extended period of time. The average proportions of polymorphic loci and genetic diversity within populations were 55.7% (range = 28.6-85.7%) and 0.213 (range = 0.114-0.317), respectively. The proportion of total genetic diversity found among populations was similar to species with equivalent life history characters (G(ST) = 0.199). No regional patterns of variation were seen. The number of putative genotypes in each population ranged from 2 to 26. Mean genotypic diversity was 0.694, ranging from 0.223 to 0.955. Such high levels of genotypic diversity indicate that local sites are often colonized by several propagules (most likely seeds) and/or that sexual reproduction occurs within populations after establishment. An excess of heterozygosity was observed in populations with few unique genets, implying that selection for highly heterozygous individuals may occur in populations of P. lobata.

Predawn disequilibrium between plant and soil water potentials in two cold-desert shrubs

Abstract Classical water relations theory predicts that predawn plant water potential should be in equilibrium with soil water potential (soil Ψw) around roots, and many interpretations of plant water status in natural populations are based on this expectation. We examined this expectation for two salt-tolerant, cold-desert shrub species in glasshouse experiments where frequent watering assured homogeneity in soil Ψw and soil-root hydraulic continuity and where NaCl controlled soil Ψw. Plant water potentials were measured with a pressure chamber (xylem Ψp) and thermocouple psychrometers (leaf Ψw). Soil Ψw was measured with in situ thermocouple psychrometers. Predawn leaf Ψw and xylem Ψp were significantly more negative than soil Ψw, for many treatments, indicating large predawn soil-plant Ψw disequilibria: up to 1.2 MPa for Chrysothamnus nauseosus (0 and 100 mm NaCl) and 1.8 MPa for Sarcobatus vermiculatus (0, 100, 300, and 600 mm NaCl). Significant nighttime canopy water loss was one mechanism contributing to predawn disequilibrium, assessed by comparison of xylem Ψp for bagged (to minimize transpiration) and unbagged canopies, and by gas exchange measurements. However, nighttime transpiration accounted for only part of the predawn disequilibrium. Other mechanisms that could act with nighttime transpiration to generate large predawn disequilibria are described and include a model of how leaf apoplastic solutes could contribute to the phenomenon. This study is among the first to conclusively document such large departures from the expectation of predawn soil-plant equilibrium for C3 shrubs, and provides a general framework for considering relative contributions of nighttime transpiration and other plant-related mechanisms to predawn disequilibrium.

Phenotypic selection on leaf water use efficiency and related ecophysiological traits for natural populations of desert sunflowers

Plant water-use efficiency (WUE) is expected to affect plant fitness and thus be under natural selection in arid habitats. Although many natural population studies have assessed plant WUE, only a few related WUE to fitness. The further determination of whether selection on WUE is direct or indirect through functionally related traits has yielded no consistent results. For natural populations of two desert annual sunflowers, Helianthus anomalus and H. deserticola, we used phenotypic selection analysis with vegetative biomass as the proxy for fitness to test (1) whether there was direct and indirect selection on WUE (carbon isotope ratio) and related traits (leaf N, area, succulence) and (2) whether direct selection was consistent with hypothesized drought/dehydration escape and avoidance strategies. There was direct selection for lower WUE in mesic and dry H. anomalus populations, consistent with dehydration escape, even though it is the longer lived of the two species. For mesic H. anomalus, direct selection favored lower WUE and higher N, suggesting that plants may be “wasting water” to increase N delivery via the transpiration stream. For the shorter lived H. deserticola in the direr habitat, there was indirect selection for lower WUE, inconsistent with drought escape. There was also direct selection for higher leaf N, succulence and leaf size. There was no direct selection for higher WUE consistent with dehydration avoidance in either species. Thus, in these natural populations of two desert dune species higher fitness was associated with some combination direct and indirect selection for lower WUE, higher leaf N and larger leaf size. Our understanding of the adaptive value of plant ecophysiological traits will benefit from further consideration of related traits such as leaf nitrogen and more tests in natural populations.

Phenotypic Differentiation between Three Ancient Hybrid Taxa and Their Parental Species

The primary requirement for a new diploid species to arise via hybridization is ecological divergence from its parental species. Ecological divergence protects the nascent hybrid species from competition with its progenitor species and may contribute to reproductive isolation. However, the means by which hybridization might facilitate the necessary adaptive transitions are poorly understood. Here, we report the results of a glasshouse experiment in which 42 morphological and ecophysiological traits were measured in three hybrid sunflower species (Helianthus anomalus, Helianthus deserticola, and Helianthus paradoxus) and their parental species (Helianthus annuus and Helianthus petiolaris). A surprisingly high proportion of traits were extreme relative to the parental species (24%, 20%, and 39% of traits in H. anomalus, H. deserticola, and H. paradoxus, respectively). Most of the extreme traits have previously been reported in the literature as adaptations to dune (H. anomalus), high‐desert (H. deserticola), or high‐salt (H. paradoxus) habitats. We propose that hybridization has contributed to ecological divergence largely via the generation of extreme traits in segregating hybrids, a commonly observed phenomenon called “transgressive segregation.”

Helianthus Nighttime Conductance and Transpiration Respond to Soil Water But Not Nutrient Availability

We investigated the response of Helianthus species nighttime conductance (gnight) and transpiration (Enight) to soil nutrient and water limitations in nine greenhouse studies. The studies primarily used wild Helianthus annuus, but also included a commercial and early domesticate of H. annuus and three additional wild species (Helianthus petiolaris Nutt., Helianthus deserticola Heiser, and Helianthus anomalus Blake). Well-watered plants of all species showed substantial gnight (0.023–0.225 mol m−2 s−1) and Enight (0.29–2.46 mmol m−2 s−1) measured as instantaneous gas exchange. Based on the potential for transpiration to increase mass flow of mobile nutrients to roots, we hypothesized that gnight and Enight would increase under limiting soil nutrients but found no evidence of responses in all six studies testing this. Based on known daytime responses to water limitation, we hypothesized that gnight and Enight would decrease when soil water availability was limited, and results from all four studies testing this supported our hypothesis. We also established that stomatal conductance at night was on average 5 times greater than cuticular conductance. Additionally, gnight and Enight varied nocturnally and across plant reproductive stages while remaining relatively constant as leaves aged. Our results further the ability to predict conditions under which nighttime water loss will be biologically significant and demonstrate that for Helianthus, gnight can be regulated.