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Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade-tolerance.

ABSTRACT: Understorey plant communities are crucial to maintain species diversity and ecosystem processes including nutrient cycling and regeneration of overstorey trees. Most studies exploring effects of elevated CO2 concentration ([CO2]) in forests have, however, been done on overstorey trees, while understorey communities received only limited attention.The hypothesis that understorey grass species differ in shade-tolerance and development dynamics, and temporally exploit different niches under elevated [CO2], was tested during the fourth year of [CO2] treatment. We assumed stimulated carbon gain by elevated [CO2] even at low light conditions in strongly shade-tolerant Luzula sylvatica, while its stimulation under elevated [CO2] in less shade-tolerant Calamagrostis arundinacea was expected only in early spring when the tree canopy is not fully developed.We found evidence supporting this hypothesis. While elevated [CO2] stimulated photosynthesis in L. sylvatica mainly in the peak of the growing season (by 55%-57% in July and August), even at low light intensities (50 µmol m-2 s-1), stimulatory effect of [CO2] in C. arundinacea was found mainly under high light intensities (200 µmol m-2 s-1) at the beginning of the growing season (increase by 171% in May) and gradually declined during the season. Elevated [CO2] also substantially stimulated leaf mass area and root-to-shoot ratio in L. sylvatica, while only insignificant increases were observed in C. arundinacea.Our physiological and morphological analyses indicate that understorey species, differing in shade-tolerance, under elevated [CO2] exploit distinct niches in light environment given by the dynamics of the tree canopy.

SUBMITTER: Holub P

PROVIDER: S-EPMC6953567 | biostudies-literature | 2019 Dec

REPOSITORIES: biostudies-literature

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Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade-tolerance.

Holub Petr P Klem Karel K Linder Sune S Urban Otmar O

Ecology and evolution 20191129 24

Understorey plant communities are crucial to maintain species diversity and ecosystem processes including nutrient cycling and regeneration of overstorey trees. Most studies exploring effects of elevated CO2 concentration ([CO2]) in forests have, however, been done on overstorey trees, while understorey communities received only limited attention.The hypothesis that understorey grass species differ in shade-tolerance and development dynamics, and temporally exploit differen ...[more]

PMID: 31938473

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Project description:Background and aimsA long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried.MethodsThe biomass distribution of juveniles (17-740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain.ResultsAn index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR(d) = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey.ConclusionsThe results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.

Project description:The carbon balance is defined here as the partitioning of daily whole-plant gross CO2 assimilation (A) in C available for growth and C required for respiration (R). A scales positively with growth irradiance and there is evidence for an irradiance dependence of R as well. Here we ask if R as a fraction of A is also irradiance dependent, whether there are systematic differences in C-balance between shade-tolerant and shade-intolerant species, and what the causes could be. Growth, gas exchange, chemical composition and leaf structure were analyzed for two shade-tolerant and three shade-intolerant herbaceous species that were hydroponically grown in a growth room at five irradiances from 20 μmol m(-2) s(-1) (1.2 mol m(-2) day(-1)) to 500 μmol m(-2) s(-1) (30 mol m(-2) day(-1)). Growth analysis showed little difference between species in unit leaf rate (dry mass increase per unit leaf area) at low irradiance, but lower rates for the shade-tolerant species at high irradiance, mainly as a result of their lower light-saturated rate of photosynthesis. This resulted in lower relative growth rates in these conditions. Daily whole-plant R scaled with A in a very tight manner, giving a remarkably constant R/A ratio of around 0.3 for all but the lowest irradiance. Although some shade-intolerant species showed tendencies toward a higher R/A and inefficiencies in terms of carbon and nitrogen investment in their leaves, no conclusive evidence was found for systematic differences in C-balance between the shade-tolerant and intolerant species at the lowest irradiance. Leaf tissue of the shade-tolerant species was characterized by high dry matter percentages, C-concentration and construction costs, which could be associated with a better defense in shade environments where leaf longevity matters. We conclude that shade-intolerant species have a competitive advantage at high irradiance due to superior potential growth rates, but that shade-tolerance is not necessarily associated with a better C-balance at low irradiance. Under those conditions tolerance to other stresses is probably more important for the performance of shade-tolerant species.

Dataset Information

Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade-tolerance.

Publications

Distinct seasonal dynamics of responses to elevated CO<sub>2</sub> in two understorey grass species differing in shade-tolerance.

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