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Seed germination and seedling emergence are key processes for population recruitment. Flooding and grazing are disturbances forming gaps that may strongly influence recruitment patterns in space and time, but their combined effects and action mechanisms have rarely been addressed. In this study we analysed the effects of microhabitat conditions associated with winter flooding and spring-summer defoliation on seed germination and seedling establishment of Paspalum dilatatum, a dominant perennial C4 grass in native grasslands of the Flooding Pampa, Argentina. The dynamics of seedling emergence from natural seed banks and buried seeds was studied in a factorial experiment with flooding and defoliation treatments applied to soil monoliths (mesocosms) collected from natural grassland. Additional laboratory experiments were applied to investigate seed germination under different combinations of temperature, light quality and simulated flooding. Seed germination and seedling emergence of P. dilatatum were promoted by flooding and high intensity defoliation. Gaps generated by flooding were maintained by high intensity defoliation exercising a synergistic effect on survival seedlings. Flooding resulted in the breaking of seed dormancy and higher germination rates associated with alternating temperature and the activation of the phytochrome system. Our results indicate that microhabitat conditions associated with the disturbances forming gaps, such as flooding and heavy grazing, synergistically promote the recruitment process of this dominant grass species.

Water supply affects seed germination and seedling establishment of shallow-rooted warm-season grasses. This may explain the difficulty of incorporating Dallisgrass (Paspalum dilatatum Poir.) into humid temperate grasslands through interseeding. We studied the effects of water availability on seed germination and seedling growth under controlled conditions to determine which step of the establishment process was most affected. In a laboratory experiment, seeds were germinated at 0, -0.25, -0.5, -0.75, and -1 MPa water availability generated with solutions of polyethylene glycol. Although both maximum rate and total germination (P < 0.05) significantly decreased with increased water stress, the speed of germination was even more sensitive. In a greenhouse experiment, variations in seedling emergence and morphological characteristics were measured in relation to water availability. Pregerminated and dry seeds were sown in pots that were irrigated at 1-, 2- , 4-, or 7-day intervals. This species showed high sensitivity to water stress during germination and early emergence. High emergence was obtained from the daily irrigation treatment. In all other treatments, where watering frequency was extended, emergence was decreased. Results suggest that rapid germination and early adventitious root growth can be obtained only with reliable availability of water. These conditions, combined with the high temperatures required for breaking seed dormancy, occur infrequently, explaining the difficulty of achieving successful establishment of Dallisgrass. Water availability during seed germination and seedling emergence is a critical factor for survival of this species.

Lolium multiflorum Lam., a winter annual grass species originating from the Mediterranean region, has become widespread in native grasslands of the flooding pampa (Argentina). The dynamics of dormancy release of L. multiflorum seeds in simulated natural conditions was studied. Dormancy was overcome by exposing imbibed seeds to mild cool temperatures (15°C or below) for brief periods (pre-chilling). After-ripening progressively released dormancy, at a higher rate in seeds that remained under the canopy (30/20°C) than those that were dry-stored (20°C). Germination was promoted by red light only when the after-ripening requirement was partially fulfilled or after pre-chilling treatment. These results provide useful knowledge for designing grazing strategies aimed at obtaining well-developed ryegrass canopy of high forage quality during winter and spring.

The aim of this study was to evaluate the possibility to use neighbour species composition to explain demographic variations in seasonal tillers in perennial grasses. Two warm-season species of the Argentina flooded pampa, Paspalum dilatatum and Sporobolus indicus, were used as models. Tiller structure and dynamics of target plants and the identity of their nearest neighbours were analyzed within a natural grassland community for an entire annual growth cycle. Canonical Correlation Analysis of tillering behaviour of target plants and neighbour species composition showed significant relationships for summer neighbourhood data. Community components affecting tiller demography were identified, and their spatial variation was described as different gradients of neighbourhood composition (NCG). NCG varied between target species, determining different spatial variation in neighbouring canopy dynamics and tiller dynamics. The tiller structure and dynamics of P. dilatatum plants were significantly correlated with the seasonal pattern of their neighbouring canopy density (NCD). Plants placed on the NCD-fluctuating extremes along NCG showed an unstable modular structure and short-lived tillers, as compared to those placed in the centre of the gradient. Relative density of the neighbouring canopy showed little variation along the NCG of S. indicus plants, which presented a tiller structure more stable and uniform than P. dilatatum plants did. Identification of NCG may be a promising approach to understanding changes in tiller dynamics of target species in relation to spatial and temporal changes in community structure.

The general objectives of this study were: (1) to investigate the importance of internal influences in regulating the tiller dynamics in natural populations of the warm-season perennial grasses Paspalum dilatatum and Sporobolus indicus, coexisting in Argentine flooded pampa, in as much as they act independently of the underlying external environment, and (2) to evaluate the extent to which interactions between internal and external factors affect the variation in tiller dynamics within such populations. Within-population variation in seasonal development of plants and tillers with different neighbour composition was studied for an annual growth cycle. Tiller survival and tillering were significantly influenced by tiller size. Tiller age influenced tiller fate, as suggested by the additive effects of age and size of tillers. These relationships varied with season and with species. Size and age of tillers showed additive effects with their neighbouring species on the tiller fate of P. dilatatum, but the effects of age and size of S. indicus changed according their neighbourhood. Tiller survival of S. indicus during the early growth season was more size-dependent when the cold-season species Poa lanigera, was the principal neighbour. Flowering and tillering probabilities were positively related through their common positive responses to tiller size. Tiller survival and recruitment between different seasons were strongly related. Independently of neighbour composition, tiller survival was generally inversely related to tiller recruitment in previous seasons. Therefore, significant density-dependent mortality of tillers was found for both species during the early summer when tiller density was expressed by basal area units.

The effects of plant density on shoot -and leaf lamina zenith angles (i.e. the angles with respect to the vertical) were investigated in Lolium multiflorum and Paspalum dilatatum plants grown in pots either outdoors or in Tillers appeared above the ligule of their subtending leaf with a small zenith angle but, in plants grown at low densities, gradually adopted a more horizontal position. In plants grown at high densities this gradual increase in shoot zenith angle with age was strongly reduced. The average shoot zenith angle at a given time was lower (i.e. tillers were more erect) at high

Casal, J. J., Sanchez, R. A. and Deregibus, V. A. 1987. Tillering responses of Lolium multiflorum plants to changes of red/far-red ratio typical of sparse canopies.—J. exp. Bot. 38: 1432-1439.