Two sides to every leaf: water and CO2 transport in hypostomatous and amphistomatous leaves.

Leaves with stomata on both upper and lower surfaces, termed amphistomatous, are relatively rare compared with hypostomatous leaves with stomata only on the lower surface. Amphistomaty occurs predominantly in fast-growing herbaceous annuals and in slow-growing perennial shrubs and trees. In this paper, we present the current understanding and hypotheses on the costs and benefits of amphistomaty related to water and CO2 transport in contrasting leaf morphologies. First, there is no evidence that amphistomatous species achieve higher stomatal densities on a projected leaf area basis than hypostomatous species, but two-sided gas exchange is less limited by boundary layer effects. Second, amphistomaty may provide a specific advantage in thick leaves by shortening the pathway for CO2 transport between the atmosphere and the chloroplasts. In thin leaves of fast-growing herbaceous annuals, in which both the adaxial and abaxial pathways are already short, amphistomaty enhances leaf-atmosphere gas-exchange capacity. Third, amphistomaty may help to optimise the leaf-interior water status for CO2 transport by reducing temperature gradients and so preventing the condensation of water that could limit CO2 diffusion. Fourth, a potential cost of amphistomaty is the need for additional investments in leaf water transport tissue to balance the water loss through the adaxial surface.

Amphistomatic leaf surfaces independently regulate gas exchange in response to variations in evaporative demand.

The occurrence of amphistomatic leaves (stomata on both surfaces) versus hypostomatic leaves (stomata limited to the lower or abaxial surface) has strong associations with environment. Amphistomy provides the advantage of higher conductance of CO2 for photosynthesis, however, unless the stomata on both leaf surfaces can be independently controlled in response to environmental cues, amphistomy may lead to inefficient gas exchange. While previous studies have found evidence that stomata can operate independently across and between surfaces of dorsiventral leaves, we investigate whether an independent stomatal response can be induced for isobilateral leaves by largely natural conditions. Here, we exposed surfaces of isobilateral, amphistomatic Eucalyptus globulus Labill. leaves to natural diurnal variation in differential evaporative demand, using leaf orientation to drive differences in irradiance and heat load on leaf surfaces. We identified preferential closure of stomata on the surface exposed to higher irradiation (and therefore evaporative demand) during the afternoon under natural conditions and similarly induced differential stomatal closure under experimental conditions in the laboratory. The differential response confirms that sufficient hydraulic isolation exists for independent stomatal response to occur between surfaces of amphistomatic, isobilateral leaves, and importantly, we show that natural conditions can induce surface-specific stomatal closure.

Improving respiration measurements with gas exchange analyzers.

Dark respiration measurements with open-flow gas exchange analyzers are often questioned for their low accuracy as their low values often reach the precision limit of the instrument. Respiration was measured in five species, two hypostomatous (Vitis Vinifera L. and Acanthus mollis) and three amphistomatous, one with similar amount of stomata in both sides (Eucalyptus citriodora) and two with different stomata density (Brassica oleracea and Vicia faba). CO2 differential (ΔCO2) increased two-fold with no change in apparent Rd, when the two leaves with higher stomatal density faced outside. These results showed a clear effect of the position of stomata on ΔCO2. Therefore, it can be concluded that leaf position is important to guarantee the improvement of respiration measurements increasing ΔCO2 without affecting the respiration results by leaf or mass units. This method will help to increase the accuracy of leaf respiration measurements using gas exchange analyzers.

Density, size and distribution of stomata in 35 rainforest tree species in Central Amazonia

Stomata are turgor-operated valves that control water loss and CO2 uptake during photosynthesis, and thereby water relation and plant biomass accumulation is closely related to stomatal functioning. The aims of this work were to document how stomata are distributed on the leaf surface and to determine if there is any significant variation in stomatal characteristics among Amazonian tree species, and finally to study the relationship between stomatal density (S D) and tree height. Thirty five trees (>17 m tall) of different species were selected. Stomatal type, density (S D), size (S S) and stomatal distribution on the leaf surface were determined using nail polish imprints taken from both leaf surfaces. Irrespective of tree species, stomata were located only on the abaxial surface (hypostomaty), with large variation in both S D and S S among species. S D ranged from 110 mm-2 in Neea altissima to 846 mm-2 in Qualea acuminata. However, in most species S D ranges between 271 and 543 mm-2, with a negative relationship between S D and S S. We also found a positive relationship between S D and tree height (r² = 0.14, p 0.01), but no correlation was found between S D and leaf thickness. The most common stomatal type was anomocytic (37%), followed by paracytic (26%) and anisocytic (11%). We conclude that in Amazonian tree species, stomatal distribution on the leaf surface is a response most likely dependent on the genetic background of every species, rather than a reaction to environmental changes, and that somehow S D is influenced by environmental factors dependent on tree height.

Estômatos são válvulas operadas a turgor que controlam a perda de água e a captura de CO2 durante a fotossíntese. Assim, as relações hídricas e o acumulo de biomassa vegetal são fortemente influenciadas pelo funcionamento estomático. Os objetivos deste trabalho foram: documentar como os estômatos estão distribuídos na superfície foliar e determinar se existe variação das características estomáticas entre espécies da Amazônia, estudar a relação entre densidade estomática (S D) e altura arbórea. Trinta e cinco árvores (>17 m de altura) de diferentes espécies foram selecionadas. Tipo de complexo estomático, S D, tamanho (S S) e distribuição na superfície foliar foram determinados utilizando impressões de ambas as superfícies foliares com esmalte incolor. Independente da espécie, os estômatos foram encontrados apenas na superfície abaxial (hipoestomatia) com ampla variação na S D e no S S entre espécies. A densidade estomática variou de 110 mm-2 em Neea altissima a 846 mm-2 em Qualea acuminata. Entretanto, a maioria das espécies apresentou S D entre 271 e 543 mm-2, com uma relação negativa entre S D e S S. Observou-se uma relação positiva entre S D e altura arbórea (r² = 0.14, p 0.01), não havendo relação entre S D e espessura foliar. Os tipos estomáticos mais comuns foram: anomocíticos (37%), seguidos de paracíticos (26%) e anisocíticos (11%). Concluiu-se que em espécies da Amazônia, a distribuição de estômatos na superfície foliar está mais relacionada a fatores genéticos de cada espécie do que a variações ambientais. Entretanto, S D é fortemente influenciada por fatores ambientais concernentes à altura da árvore.

Density, size and distribution of stomata in 35 rainforest tree species in Central Amazonia / Densidade, tamanho e distribuição estomática em 35 espécies de árvores na Amazônia Central

Stomata are turgor-operated valves that control water loss and CO2 uptake during photosynthesis, and thereby water relation and plant biomass accumulation is closely related to stomatal functioning. The aims of this work were to document how stomata are distributed on the leaf surface and to determine if there is any significant variation in stomatal characteristics among Amazonian tree species, and finally to study the relationship between stomatal density (S D) and tree height. Thirty five trees (>17 m tall) of different species were selected. Stomatal type, density (S D), size (S S) and stomatal distribution on the leaf surface were determined using nail polish imprints taken from both leaf surfaces. Irrespective of tree species, stomata were located only on the abaxial surface (hypostomaty), with large variation in both S D and S S among species. S D ranged from 110 mm-2 in Neea altissima to 846 mm-2 in Qualea acuminata. However, in most species S D ranges between 271 and 543 mm-2, with a negative relationship between S D and S S. We also found a positive relationship between S D and tree height (r² = 0.14, p < 0.01), but no correlation was found between S D and leaf thickness. The most common stomatal type was anomocytic (37%), followed by paracytic (26%) and anisocytic (11%). We conclude that in Amazonian tree species, stomatal distribution on the leaf surface is a response most likely dependent on the genetic background of every species, rather than a reaction to environmental changes, and that somehow S D is influenced by environmental factors dependent on tree height.

Estômatos são válvulas operadas a turgor que controlam a perda de água e a captura de CO2 durante a fotossíntese. Assim, as relações hídricas e o acumulo de biomassa vegetal são fortemente influenciadas pelo funcionamento estomático. Os objetivos deste trabalho foram: documentar como os estômatos estão distribuídos na superfície foliar e determinar se existe variação das características estomáticas entre espécies da Amazônia, estudar a relação entre densidade estomática (S D) e altura arbórea. Trinta e cinco árvores (>17 m de altura) de diferentes espécies foram selecionadas. Tipo de complexo estomático, S D, tamanho (S S) e distribuição na superfície foliar foram determinados utilizando impressões de ambas as superfícies foliares com esmalte incolor. Independente da espécie, os estômatos foram encontrados apenas na superfície abaxial (hipoestomatia) com ampla variação na S D e no S S entre espécies. A densidade estomática variou de 110 mm-2 em Neea altissima a 846 mm-2 em Qualea acuminata. Entretanto, a maioria das espécies apresentou S D entre 271 e 543 mm-2, com uma relação negativa entre S D e S S. Observou-se uma relação positiva entre S D e altura arbórea (r² = 0.14, p < 0.01), não havendo relação entre S D e espessura foliar. Os tipos estomáticos mais comuns foram: anomocíticos (37%), seguidos de paracíticos (26%) e anisocíticos (11%). Concluiu-se que em espécies da Amazônia, a distribuição de estômatos na superfície foliar está mais relacionada a fatores genéticos de cada espécie do que a variações ambientais. Entretanto, S D é fortemente influenciada por fatores ambientais concernentes à altura da árvore.