RESUMEN
Stomatal responses to environmental signals differ substantially between ferns and angiosperms. However, the mechanisms that lead to such different responses remain unclear. Here we investigated the extent to which leaf metabolism contributes to coordinate the differential stomatal behaviour among ferns and angiosperms. Stomata from all species were responsive to light and CO2 transitions. However, fern stomatal responses were slower and minor in both absolute and relative terms. Angiosperms have higher stomatal density, but this is not correlated with speed of stomatal closure. The metabolic responses throughout the diel course and under different CO2 conditions differ substantially among ferns and angiosperms. Higher sucrose content and an increased sucrose-to-malate ratio during high CO2 -induced stomatal closure was observed in angiosperms compared to ferns. Furthermore, the speed of stomatal closure was positively and negatively correlated with sugars and organic acids, respectively, suggesting that the balance between sugars and organic acids aids in explaining the faster stomatal responses of angiosperms. Our results suggest that mesophyll-derived metabolic signals, especially those associated with sucrose and malate, may also be important to modulate the differential stomatal behaviour between ferns and angiosperms, providing important new information that helps in understanding the metabolism-mediated mechanisms regulating stomatal movements across land plant evolution.
Asunto(s)
Dióxido de Carbono/metabolismo , Helechos/fisiología , Luz , Magnoliopsida/fisiología , Malatos/metabolismo , Estomas de Plantas/metabolismo , Estomas de Plantas/efectos de la radiación , Sacarosa/metabolismo , Análisis Discriminante , Helechos/efectos de la radiación , Análisis de los Mínimos Cuadrados , Magnoliopsida/efectos de la radiación , Metaboloma/efectos de la radiación , Fotosíntesis/efectos de la radiación , Análisis de Componente PrincipalRESUMEN
Background and Aims: The greater diversity of plant clades in the Neotropics compared to their relatives in Africa is a pervasive pattern in biogeography. To better understand the causes of this imbalance, we studied the diversification dynamics of the monocot family Velloziaceae. In addition to being conspicuously richer in the Neotropics compared to the Palaeotropics, many species of Velloziaceae exhibit extreme desiccation tolerance (i.e. 'resurrection' behaviour), and other ecological specializations to life on rocky outcrops, poor sandy soils, open vegetation and seasonally dry climates. Velloziaceae is also ecologically dominant in the campos rupestres, a habitat having exceptionally high plant diversity and endemism in Brazil. Methods: We reconstructed a densely sampled time-calibrated molecular phylogeny and used state-dependent and state-independent models to estimate rates of lineage diversification in relation to continent-scale geographical occurrence and functional traits associated with desiccation tolerance and water storage capacity. Key Results: Independent shifts to faster diversification occurred within two Neotropical lineages, Vellozia and Barbacenia. The Vellozia radiation was associated with the presence of conspicuous aerial stems, and was followed by decreasing diversification rates during the Oligocene, a time of rising global temperatures and expanding open areas around the world. The Barbacenia radiation was faster and more recent, occurring during the cooling conditions of the Miocene, and associated with the acquisition of aquiferous parenchyma on the leaves. Conclusions: High species richness of Velloziaceae in South America has been driven by faster diversification in lineages predominantly occurring in the campos rupestres, putatively by the evolution of adaptive strategies in response to independent climatic events. The radiation of Vellozia in particular might have played a key role in the assembly of the campos rupestres vegetation.
Asunto(s)
Biodiversidad , Magnoliopsida/genética , Adaptación Fisiológica , Evolución Biológica , Brasil , Desecación , Ecosistema , Magnoliopsida/fisiología , Magnoliopsida/efectos de la radiación , Fenotipo , Filogenia , Tallos de la Planta/genética , Tallos de la Planta/fisiología , Tallos de la Planta/efectos de la radiaciónRESUMEN
The relationship between the phytohormones, gibberellin (GA) and abscisic acid (ABA) and light and temperature on seed germination is still not well understood. We aimed to investigate the role of the ABA and GA on seed germination of Vellozia caruncularis, V. intermedia and V. alutacea in response to light/dark conditions on different temperature. Seeds were incubated in GA (GA3 or GA4 ) or ABA and their respective biosynthesis inhibitors (paclobutrazol - PAC, and fluridone - FLU) solutions at two contrasting temperatures (25 and 40 °C). Furthermore, endogenous concentrations of active GAs and those of ABA were measured in seeds of V. intermedia and V. alutacea during imbibition/germination. Exogenous ABA inhibited the germination of Vellozia species under all conditions tested. GA, FLU and FLU + GA3 stimulated germination in the dark at 25 °C (GA4 being more effective than GA3 ). PAC reduced seed germination in V. caruncularis and V. alutacea, but did not affect germination of V. intermedia at 40 °C either under light or dark conditions. During imbibition in the dark, levels of active GAs decreased in the seeds of V. intermedia, but were not altered in those of V. alutacea. Incubation at 40 °C decreased ABA levels during imbibition in both V. caruncularis and V. alutacea. We conclude that the seeds of Vellozia species studied here require light or high temperature to germinate and ABA has a major role in the regulation of Vellozia seed germination in response to light and temperature.
Asunto(s)
Ácido Abscísico/farmacología , Regulación de la Expresión Génica de las Plantas , Germinación/efectos de los fármacos , Giberelinas/farmacología , Magnoliopsida/efectos de los fármacos , Reguladores del Crecimiento de las Plantas/farmacología , Ácido Abscísico/antagonistas & inhibidores , Oscuridad , Giberelinas/antagonistas & inhibidores , Calor , Luz , Magnoliopsida/fisiología , Magnoliopsida/efectos de la radiación , Reguladores del Crecimiento de las Plantas/antagonistas & inhibidores , Piridonas/farmacología , Semillas/efectos de los fármacos , Semillas/fisiología , Semillas/efectos de la radiación , Triazoles/farmacologíaRESUMEN
Several studies across species have linked leaf functional traits with shade tolerance. Because evolution by natural selection occurs within populations, in order to explain those interspecific patterns it is crucial to examine variation of traits associated with shade tolerance and plant fitness at an intraspecific scale. In a southern temperate rainforest, two climbing plant species coexist but differ in shade tolerance. Whereas Luzuriaga radicans is most abundant in the shaded understory, L. polyphylla typically occurs in intermediate light environments. We carried out an intraspecific approach to test the hypothesis of differential selection patterns in relation to shade tolerance in these congeneric species. The probability of showing reproductive structures increased with specific leaf area (SLA) in L. polyphylla, and decreased with dark respiration in L. radicans. When reproductive output of fertile individuals was the fitness variable, we detected positive directional selection on SLA in L. polyphylla, and negative directional selection on dark respiration and positive directional selection on leaf size in L. radicans. Total light radiation differed between the microsites where the Luzuriaga species were sampled in the old-growth forest understory. Accordingly, L. radicans had a lower minimum light requirement and showed fertile individuals in darker microsites. L. radicans showed lower dark respiration, higher chlorophyll content, and greater leaf size and SLA than L. polyphylla. Results suggest that in more shade-tolerant species, established in the darker microsites, selection would favor functional traits minimizing carbon losses, while in less shade-tolerant species, plants displaying leaf traits enhancing light capture would be selected.
Asunto(s)
Magnoliopsida/fisiología , Selección Genética , Luz Solar , Chile , Magnoliopsida/metabolismo , Magnoliopsida/efectos de la radiación , Análisis Multivariante , Fenotipo , Fotosíntesis , Desarrollo de la Planta/efectos de la radiación , Hojas de la Planta/metabolismo , Hojas de la Planta/fisiología , Hojas de la Planta/efectos de la radiación , Análisis de Regresión , Especificidad de la Especie , Clima TropicalRESUMEN
Foliar anatomy and morphology are strongly related to physiological performance; therefore, phenotypic plasticity in leaves to variations in environmental conditions, such as irradiance and soil moisture availability, can be related to growth rate and survivorship, mainly during critical growth phases, such as establishment. The aim of this work was to analyze changes in the foliar internal anatomy (tissue proportions and cell dimensions) and external morphology (leaf length, width and area) of Nothofagus pumilio (Poepp. et Endl.) Krasser seedlings growing in a greenhouse under controlled irradiance (three levels) and soil moisture (two levels) during one growing season (measured three times), and to relate them to physiological traits. Three irradiance levels (4, 26 and 64% of the natural incident light) and two soil moisture levels (40 and 80% soil capacity) were evaluated during November, January and March. Internal foliar anatomy of seedlings was analyzed using digital photographs of histological cuttings, while leaf gross morphology was measured using digital calipers and image analysis software. Most internal anatomical variables presented significant differences under different irradiance levels during the growing season, but differences were not detected between soil moisture levels. Palisade parenchyma was the tissue most sensitive to irradiance levels, and high irradiance levels (64% natural incident light) produced greater values in most of the internal anatomical variables than lower irradiance levels (4-24% natural incident light). Complementarily, larger leaves were observed in medium and low irradiance levels, as well as under low soil moisture levels (40% soil capacity). The relationship of main results with some eco-physiological traits was discussed. Foliar internal anatomical and external morphological plasticity allows quick acclimation of seedlings to environmental changes (e.g., during harvesting). These results can be used to propose new forest practices that consider soil moisture and light availability changes to maintain high physiological performance of seedlings.
Asunto(s)
Magnoliopsida/anatomía & histología , Magnoliopsida/crecimiento & desarrollo , Hojas de la Planta/anatomía & histología , Árboles/anatomía & histología , Árboles/crecimiento & desarrollo , Aclimatación , Ambiente , Magnoliopsida/fisiología , Magnoliopsida/efectos de la radiación , Fotosíntesis , Hojas de la Planta/crecimiento & desarrollo , Hojas de la Planta/fisiología , Hojas de la Planta/efectos de la radiación , Distribución Aleatoria , Estaciones del Año , Plantones/anatomía & histología , Plantones/crecimiento & desarrollo , Plantones/fisiología , Plantones/efectos de la radiación , Suelo/química , Luz Solar , Árboles/fisiología , Árboles/efectos de la radiaciónRESUMEN
Nothofagus dombeyi (Mirb.) Blume and Nothofagus nitida (Phil.) Krasser, two evergreens in the South Chilean forest, regenerate in open habitats and under the canopy, respectively. Both overtop the forest canopy when they are in the adult stage, suggesting that their photoprotective mechanisms differ in ontogenetic dynamics. We postulated that N. nitida, a shade-tolerant species increases its capacity to tolerate photoinhibitory conditions (low temperature and high irradiance) by thermal energy dissipation of excess energy during its transition from the seedling to the adult stage, whereas N. dombeyi, a shade-intolerant species, maintains a high capacity for photoprotection by thermal energy dissipation from the seedling to the adult stage. To test this hypothesis, the main photoprotective mechanisms in plants - the fast- and slow-relaxing components of thermal energy dissipation (NPQ, non-photochemical quenching) NPQ(F) and NPQ(S), respectively, and state transitions - were studied in seedlings and adults of both species grown in their natural habitats and in a common garden. In adults, NPQ(F) and NPQ(S) did not differ between species and seasons. The greatest differences in these parameters were observed in seedlings. The xanthophyll cycle was more active in N. dombeyi seedlings than in N. nitida seedlings at low temperature and high irradiance, consistent with a higher NPQ(F) in N. dombeyi. Under all study conditions, N. nitida seedlings had higher NPQ(S) than N. dombeyi seedlings. The state transition capability was higher in N. nitida seedlings than in N. dombeyi seedlings. Therefore, although (shade-intolerant) N. dombeyi was able to thermally dissipate the excess absorbed energy, under natural conditions its photochemical energy quenching was efficient in both developmental stages, decreasing its need for thermal dissipation. In contrast, the seedlings of N. nitida were more sensitive to photoinhibition than the adult trees, suggesting a change from shade-grown to sun-exposed phenotype from the seedling to the adult stage. These results help to explain the differences in the regeneration patterns of N. nitida and N. dombeyi and the presence of N. nitida adult stage in the upper canopy.
Asunto(s)
Magnoliopsida/crecimiento & desarrollo , Estaciones del Año , Luz Solar , Magnoliopsida/metabolismo , Magnoliopsida/efectos de la radiación , Pigmentos Biológicos/metabolismo , Plantones/crecimiento & desarrollo , Plantones/metabolismo , Plantones/efectos de la radiación , Especificidad de la Especie , Temperatura , Xantófilas/metabolismoRESUMEN
The extent to which Crassulacean acid metabolism (CAM) plant delta(13)C values provide an index of the proportions of CO(2) fixed during daytime and nighttime was assessed. Shoots of seven CAM species (Aloe vera, Hylocereus monocanthus, Kalanchoe beharensis, Kalanchoe daigremontiana, Kalanchoe pinnata, Vanilla pauciflora, and Xerosicyos danguyi) and two C(3) species (teak [Tectona grandis] and Clusia sp.) were grown in a cuvette, and net CO(2) exchange was monitored for up to 51 d. In species exhibiting net dark CO(2) fixation, between 14% and 73.3% of the carbon gain occurred in the dark. delta(13)C values of tissues formed inside the cuvette ranged between -28.7 per thousand and -11.6 per thousand, and correlated linearly with the percentages of carbon gained in the light and in the dark. The delta(13)C values for new biomass obtained solely during the dark and light were estimated as -8.7 per thousand and -26.9 per thousand, respectively. For each 10% contribution of dark CO(2) fixation integrated over the entire experiment, the delta(13)C content of the tissue was, thus, approximately 1.8 per thousand less negative. Extrapolation of the observations to plants previously surveyed under natural conditions suggests that the most commonly expressed version of CAM in the field, "the typical CAM plant," involves plants that gain about 71% to 77% of their carbon by dark fixation, and that the isotopic signals of plants that obtain one-third or less of their carbon in the dark may be confused with C(3) plants when identified on the basis of carbon isotope content alone.
Asunto(s)
Dióxido de Carbono/metabolismo , Magnoliopsida/metabolismo , Fotosíntesis/fisiología , Proteínas del Complejo del Centro de Reacción Fotosintética/metabolismo , Isótopos de Carbono , Oscuridad , Luz , Magnoliopsida/clasificación , Magnoliopsida/efectos de la radiación , Brotes de la Planta/crecimiento & desarrollo , Brotes de la Planta/metabolismo , Factores de TiempoRESUMEN
The primary motivation behind the considerable effort in studying stratospheric ozone depletion is the potential for biological consequences of increased solar UVB (280-315 nm) radiation. Yet, direct links between ozone depletion and biological impacts have been established only for organisms of Antarctic waters under the influence of the ozone "hole;" no direct evidence exists that ozone-related variations in UVB affect ecosystems of temperate latitudes. Indeed, calculations based on laboratory studies with plants suggest that the biological impact of ozone depletion (measured by the formation of cyclobutane pyrimidine dimers in DNA) is likely to be less marked than previously thought, because UVA quanta (315-400 nm) may also cause significant damage, and UVA is unaffected by ozone depletion. Herein, we show that the temperate ecosystems of southern South America have been subjected to increasingly high levels of ozone depletion during the last decade. We found that in the spring of 1997, despite frequent cloud cover, the passages of the ozone hole over Tierra del Fuego (55 degrees S) caused concomitant increases in solar UV and that the enhanced ground-level UV led to significant increases in DNA damage in the native plant Gunnera magellanica. The fluctuations in solar UV explained a large proportion of the variation in DNA damage (up to 68%), particularly when the solar UV was weighted for biological effectiveness according to action spectra that assume a sharp decline in quantum efficiency with increasing wavelength from the UVB into the UVA regions of the spectrum.