UNIVERSIDAD NACIONAL DEL SUR

Tesis de Doctor en Biología

Ecofisiología de plantas del sotobosque de Nothofagus pumilio: efectos de la apertura del dosel

Luciano Javier Selzer

Bahía Blanca Argentina 2013 Resumen La apertura del dosel del bosque generada por disturbios antrópicos o naturales provoca una alteración en las condiciones ambientales del sistema ecológico, modificando la cantidad y calidad de luz, las precipitaciones, y la temperatura. Los estudios realizados en este trabajo de tesis determinaron algunos de los cambios morfo-fisiológicos que permiten el crecimiento, desarrollo y reproducción de tres especies herbáceas, Osmorhiza depauperata, Phleum alpinum y Poa pratensis, en los ambientes boscosos modificados. Estas especies aumentan su cobertura luego de la cosecha forestal, y podrían evitar la erosión del suelo, y facilitar o competir con las plántulas que podrían contribuir a regenerar el bosque. En el bosque aprovechado forestalmente mediante la prescripción de retención variable se identificaron tres situaciones donde crecieron las plantas de las especies herbáceas: dentro de la retención agregada (RA), en el borde de la retención dispersa con influencia del agregado (BRA), y en la retención dispersa (RD). Además, se seleccionaron plantas del bosque primario (BP). Por otro lado, P. alpinum y P. pratensis fueron expuestas en el invernáculo a tres niveles de radiación incidente (I4: 4%; I26: 26%; I64: 64% de la luz solar incidente fuera del bosque en los diferentes ambientes), y dos niveles de contenido de humedad del suelo (M30: 30- 50% y M60: 60-80% de capacidad de campo). Las intensidades lumínicas medidas siguieron el orden: RD > BRA > RA > BP. Las plantas de todas las especies modificaron su morfo-fisiología en los distintos ambientes. Estos cambios incluyeron modificaciones en el largo de las hojas, y por consiguiente su área foliar, aumentando en los ambientes con menor radiación (BP y RA). Asimismo, en estos ambientes disminuyeron el macollaje o la cantidad de tallos. Las plantas también incrementaron la densidad de longitud de raíces en respuesta al incremento en la disponibilidad de luz. Las diferencias entre especies se podrían deber a sus hábitos de crecimiento y a la proporción de raíces finas que producen. En O. depauperata el grado de infección por micorrizas vesículo-arbusculares fue mayor que en las especies de gramíneas P. alpinum y P. pratensis. Para todas las especies se encontró que el grado de infección tendió a incrementarse al aumentar la luz, aunque solo se encontró que fue significativamente menor en BP que en el resto de los ambientes. Todas las especies produjeron mayor cantidad de biomasa por unidad de superficie

iv a medida que aumentó la irradiación: el orden de las especies estuvo inversamente relacionado con el tamaño del área basal, fue mayor en O. depauperata que en P. alpinum y P. pratensis. La cantidad de inflorescencias por unidad de área basal siguió la misma tendencia que la biomasa, y no se encontraron cambios en la proporción de inflorescencias por unidad de biomasa excepto en P. alpinum. En esta especie, esta variable fue menor en BP que en el resto de los ambientes. En P. pratensis el esfuerzo reproductivo fue mayor que en O. depauperata y P. alpinum. En invernáculo se demostró que la variable más importante que modificó el crecimiento de estas plantas fue la luz. El crecimiento disminuyó considerablemente en I4. Al disminuir la irradiación aumentó la proporción de área foliar, mediante un aumento del área foliar específica y un aumento de la proporción de biomasa foliar. Al mismo tiempo, aumentó la proporción de biomasa de tallo, y disminuyó la proporción de biomasa radical. Como resultado de estos cambios, las plantas en I26 lograron mantener una tasa de crecimiento relativa similar o superior a la de I64; sin embargo, dichos cambios no fueron suficientes para mantener la tasa de crecimiento relativa en I4. Asimismo, las especies mostraron cambios típicos a nivel fisiológico: al aumentar la irradiación aumentaron la tasa máxima de fotosíntesis y el punto de compensación lumínico. Se encontraron variaciones temporales que afectaron estos valores. Al aumentar la irradiación disminuyó el contenido de clorofila tanto por unidad de área como de peso fresco, y aumentaron las relaciones clorofila a/b y carotenoides/clorofila. Todos estos resultados indican que O. depauperata, P. alpinum y P. pratensis son capaces de aclimatarse a varias situaciones ambientales. Sin embargo, O. depauperata mostró una mejor capacidad de aclimatación a niveles de irradiación muy bajos, mientras que las especies de gramíneas apenas lograron sobrevivir dado que su crecimiento fue muy lento.

Palabras Claves: luz-sombra, aclimatación, Osmorhiza depauperata, Phleum alpinum, Poa pratensis.

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Abstract Canopy openings, whether they come from anthropic or natural disturbances, might change the environmental conditions of the ecological system, modifying light quantity and quality, precipitation and temperature. Some morpho- physiological changes that determine growth, development and reproduction of the three study herbaceous species, Osmorhiza depauperata, Phleum alpinum, Poa pratensis, were evaluated in the forest, modified environments. These species increase their coverage after forest harvesting, and could prevent soil erosion, and either facilitate or compete with seedlings that could contribute to regenerate the forest. Three environments where the herbaceous species often grow were identified in a forest previously harvested following the variable retention system. They were: within the aggregate retention (RA); on the edge of the dispersed retention under the influence of the aggregate (BRA), and dispersed retention (RD). Plants were also selected from a primary forest (BP). Furthermore, P. alpinum and P. pratensis were exposed to three levels of incident radiation (I4: 4%; I26: 26%, or I64: 64% of ambient sunlight outside the forest in the various environments) and two levels of soil moisture content (M30: 30-50% or M60: 60-80% of field capacity) under greenhouse conditions. Measured light intensities showed the following order: RD > BRA > RA> BP. Plants of all species modified their morpho-physiology in the different environments. These changes included variations in blade length, and subsequently in leaf area, increasing in the environments with lower radiation (BP and RA). Also, tillering and stem numbers decreased in these environments. Plant root length density increased as light availability also increased. Species differences could be due to their growth habits and the proportion of fine roots that they produce. Vesicular-arbuscular mycorrhizae colonization was greater in O. depauperata than in P. alpinum and P. pratensis. It also tended to decrease as light availability decreased for all species. However, it was only significantly lower in BP than in the remaining environments. Increased light availability determined a greater biomass production per unit surface area in all species: the species order was inversely related to size of the basal area. It was greater in O. depauperata than in P. alpinum and P. pratensis. The number of inflorescences per unit basal area followed the same trend as biomass and no changes were found in the proportion of

vi inflorescences per unit biomass, except in P. alpinum. In this species, this variable was lower in BP than in the other environments. Reproductive effort was greater in P. pratensis than in O. depauperata and P. alpinum. Greenhouse experiments showed that light was the major variable that modified growth of the study species. Growth decreased significantly in I4. Increases in specific leaf area and the proportion of leaf biomass determined an increased proportion of leaf area under lower light availability. At the same time, stem biomass proportion increased and that of root biomass decreased. As a result of these changes, plant relative growth rate was similar or greater in I26 than in I64; however, they were not enough to maintain plant relative growth rate in I4. Also, the species showed typical physiological changes: maximum photosynthetic rate and light compensation point increased as light availability also increased. Values for these variables varied with time. Chlorophyll content per unit surface and fresh weight decreased, and chlorophyll a/b and carotenoid/chlorophyll ratios increased, as light availability increased. The results obtained in this study indicate that O. depauperata, P. alpinum and P. pratensis are able to acclimate to various environmental situations. However, O. depauperata showed a better acclimation capacity than the other species to very low levels of light availability, while the grass species just were able to survive under these conditions because of their very slow growth.

Keywords: light-shade, acclimation, Osmorhiza depauperata, Phleum alpinum, Poa pratensis.

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