We present a meta-analysis of plant responses to fertilization experiments conducted in lowland, species-rich, tropical forests. We also update a key result and present the first species-level analyses of tree growth rates for a 15-yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community-level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low-P soils would respond most slowly. The data did not support this prediction, species-level tree growth responses to P addition were unrelated to species-level soil P associations. The meta-analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta-analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta-analysis found no support for the long-standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species-rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient-poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected.
Summary1. Drought-induced mortality and regional dieback of woody vegetation are reported from numerous locations around the world. Yet within any one site, predicting which species are most likely to survive global change-type drought is a challenge. 2. We studied the diversity of drought survival traits of a community of 15 woody plant species in a desert-chaparral ecotone. The vegetation was a mix of chaparral and desert shrubs, as well as endemic species that only occur along this margin. This vegetation boundary has large potential for drought-induced mortality because nearly all species are at the edge of their range. 3. Drought survival traits studied were vulnerability to drought-induced xylem cavitation, sapwood capacitance, deciduousness, photosynthetic stems, deep roots, photosynthetic responses to leaf water potential and hydraulic architecture. Drought survival strategies were evaluated as combinations of traits that could be effective in dealing with drought. 4. The large variation in seasonal predawn water potential of leaves and stem xylem ranged from À6Á82 to À0Á29 MPa and À6Á92 to À0Á27 MPa, respectively. The water potential at which photosynthesis ceases ranged from À9Á42 to À3Á44 MPa. Architecture was a determinant of hydraulic traits, with species supporting large leaf area per sapwood area exhibiting high rates of water transport, but also xylem that is vulnerable to drought-induced cavitation. Species with more negative midday leaf water potential during the growing season also showed access to deeper water sources based on hydrogen isotope analysis. 5. Drought survival mechanisms comprised of drought deciduousness, photosynthetic stems, tolerance of low minimum seasonal tissue water potential and vulnerability to drought-induced xylem cavitation thus varied orthogonally among species, and promote a diverse array of drought survival strategies in an arid ecosystem of considerable floristic complexity.
bstract. Lianas are a prominent growth form in tropical forests, and there is compelling evidence that they are increasing in abundance throughout the Neotropics. While recent evidence shows that soil resources limit tree growth even in deep shade, the degree to which soil resources limit lianas in forest understories, where they coexist with trees for decades, remains unknown. Regardless, the physiological underpinnings of soil resource limitation in deeply shaded tropical habitats remain largely unexplored for either trees or lianas. Theory predicts that lianas should be more limited by soil resources than trees because they occupy the quick-return end of the ''leaf economic spectrum,'' characterized by high rates of photosynthesis, high specific leaf area, short leaf life span, affinity to high-nutrient sites, and greater foliar nutrient concentrations. To address these issues, we asked whether soil resources (nitrogen, phosphorus, and potassium), alone or in combination, applied experimentally for more than a decade would cause significant changes in the morphology or physiology of tree and liana seedlings in a lowland tropical forest. We found evidence for the first time that phosphorus limits the photosynthetic performance of both trees and lianas in deeply shaded understory habitats. More importantly, lianas always showed significantly greater photosynthetic capacity, quenching, and saturating light levels compared to trees across all treatments. We found little evidence for nutrient 3 growth form interactions, indicating that lianas were not disproportionately favored in nutrient-rich habitats. Tree and liana seedlings differed markedly for six key morphological traits, demonstrating that architectural differences occurred very early in ontogeny prior to lianas finding a trellis (all seedlings were self-supporting). Overall, our results do not support nutrient loading as a mechanism of increasing liana abundance in the Neotropics. Rather, our finding that lianas always outperform trees, in terms of photosynthetic processes and under contrasting rates of resource supply of macronutrients, will allow lianas to increase in abundance if disturbance and tree turnover rates are increasing in Neotropical forests as has been suggested.
•Premise of the Study: Plant water status during flowering is important for plant reproduction, but the physiology of floral water use is not well understood. We investigated plant water status in relation to leaf and floral physiology in naturally occurring individuals of a semiarid shrub, Salvia mellifera E. Greene.•Methods: We measured stomatal (gs) and corolla (gc) conductance to water vapor, transpiration from leaves (Eleaf) and corollas (Ecorolla), leaf‐specific hydraulic conductance (KH), bulk shoot water potential (Ψshoot), and shoot water content on irrigated and control plants to analyze whether water was limiting to leaf and floral water use.•Key Results: Experimental irrigation caused a 203% increase in soil moisture content, a 20% increase in predawn Ψshoot, a 29% increase in midday Ψshoot, and a 92% increase in KH. Floral and leaf gas exchange did not respond significantly to water addition, indicating that rates were at seasonal maxima and not limited by water availability. Total daily water use by corollas was ∼20% of total shoot water use. There were no significant differences in total daily shoot water use with water addition. Mean shoot water content (5.07 g) was close to mean daily shoot water use (6.71 g), indicating that the equivalent of total shoot water content turned over every 0.76 d.•Conclusions: These results demonstrate that although irrigation improved whole‐plant hydraulic conductance, gas exchange was not limited by water availability. Additionally, the high water use of flowers in this species might limit future flowering and reproductive success during dry years.
Nitrogen (N) deposition in heavily polluted southern Californian shrublands is estimated to be 20-45 kg N x ha(-1) x yr(-1), but more exposed locales can receive as much as 145 kg N x ha(-1) x yr(-1). This large anthropogenic N input has the capacity to alter the composition of plant communities. We conducted N-fertilization experiments in chaparral and coastal sage scrub (CSS) stands over a five-year period to test the hypothesis that plant community composition would change in response to dry-season N addition because of an increase in the relative abundance of herbaceous plant species. Our results indicate that dry-season addition of N significantly altered the community composition of CSS but not chaparral. Contrary to our original hypothesis, changes in community composition were due to changes in the relative abundance of dominant shrubs and not herbaceous plant species. Given that community-level responses to changes in resource availability may take years to decades in order to fully materialize, our results suggest that continued dry-season input of N will cause even larger changes in community composition over time. These results have implications for plant species composition and diversity of mediterranean-type shrublands as N deposition increases with population growth and fossil-fuel use.
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