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树木密度第二部分
实验一 (实验组:spring时期播种a patchy distribution of bluegrass + a random distribution of 四分之一bluegrass数量的groundsel; ) (控制组:spring时期播种a random distribution of bluegrass+ a random distribution of 四分之一bluegrass数量的groundsel;) These results did not arise from neighborhood competition. Regardless of the distribution of the bluegrass, the spring generation of groundsel produced essentially the same number of seeds. That is, competition within the spring generation did not generate the differences that appeared in the fall. Pacala and Silander obtained similar results when exploring how patchiness affected the growth rates in a competitive system of velvetleaf and pigweed. (实验一结果:fall时期发现实验组的groundsel数量超出控制组4倍多.However, 这些结果不能证明理论1,因为spring时期实验组和控制组的groundsel数量差不多) On the other hand, gaps did affect the groundsel’s growth. In the plots, a large fraction of all the surviving groundsel seedlings grew in areas of low bluegrass density. So patchy plots promoted the growth of groundsel populations by providing a greater number of gaps. (相反,理论2在发挥作用,大部分fall时期发芽的groundsel是在bluegrass密度较低的地区) The difference in groundsel’s success in the spring and fall generations suggested that the bluegrass litterdead blades and roots from previous generations—could be a factor. To test that hypothesis, I repeated the above experiment with the patchy and randomly distributed bluegrass but removed the dead bluegrass from half of the plots and left the bluegrass litter intact in the other half. If litter drives the effect of spatial patterning on groundsel, then removing the litter should remove the effect, which is exactly what I found. With the litter intact, groundsel grew better where the bluegrass distribution was patchy; but with the litter removed, the groundsel grew about the same regardless of the distribution of bluegrass. This experiment indicated clearly that competition between generations, not within them, governs the dynamics of the groundsel-bluegrass system and explains the importance of the spatial pattern. (spring时期的bluegrass的残留叶子和根茎可能是一个重要因素——引出实验二) (实验组/控制组:在一半的试验种植地中,remove掉bluegrass的残留物,在另一半中,保持残留物完好无损;其他条件与实验一相同) (实验逻辑:如果隔代植物的残留物是影响spatial patterning的因素的话,那么remove掉残留物应该使这种效果消失,而此次实验确实证实了) (实验结果:在残留物完好的土地中,groundsel在patchy的分布下比在random的分布下长的更好;在残留物去除的土地中,groundsel在2种分布下长的差不多;证明理论2,而非理论1,的重要性) A series of greenhouse experiments revealed that the effect of bluegrass litter comes from the dead blades above ground. The presence of grass roots or chemicals that might have leached from the bluegrass did not affect the germination or survival of groundsel seedlings. Instead, litter inhibits groundsel seedlings, because emerging seedlings get trapped by the litter above them. The seedlings cannot penetrate the litter, which prevents them from capturing light or growing, and they die. This structural inhibition between bluegrass litter and groundsel seedlings provides the crucial competitive interaction. In addition, litter generates little trouble for the relatively slender morphology of a bluegrass seedling. These investigations illustrate that the spatial pattern of bluegrass produces large effects on the success of the competitively inferior groundsel, and that the mechanism involves gap colonization, or interactions between generations. That conclusion has several additional implications. First, the interaction between groundsel and bluegrass includes a time lag-earlier generations affecting later ones. A variety of simple mathematical models illustrate that biological systems with time lags tend to have relatively more complex dynamics than systems without time lags. The second implication involves succession. As succession proceeds, a system’s litter accumulates, which can shift the competitive balance from litter-intolerant species to litter-tolerant ones. In that way, litter can qualitatively alter the outcome of competition. Again, models support such a conclusion, showing that bluegrass should dominate whenever litter accumulates, and that groundsel dominates if the litter decomposes quickly. Modeling Trade-offs These small-scale experiments showed that groundsel grows more successfully in a patchy plot. In the simplest terms, one might say that greater amounts of bare ground favor groundsel, because the plant requires such gaps for establishment. Then one might ask: Given a particular amount of bare ground, how does its spatial distribution influence the success of invading weeds? I approached that question in collaboration with Jonathan Newman of Southern Illinois University and Ernesto Floresroux, then of the University of Chicago. We performed experiments on a somewhat larger spatial scale-over a few meters-where we tried to determine how the dispersion of gaps influences how fast groundsel progresses through a field. These experiments reveal the community-level repercussions of between-generation competition. We approached the effects of gap dispersion with a simple experiment. For each experimental plot in a field of ryegrass, we created six transects that were oriented like spokes on a wheel. On each transect, we created artificial gaps that covered one of three areas: 25, 225 or 900 square centimeters. To control the total amount of gap in a given transect, we created fewer large gaps than small ones. We distributed the gaps either uniformly or randomly, based on the distance between them. By analogy with the small-scale experiments described earlier, a large variance in the intergap distance corresponds to a patchy distribution, and equal intergap distances correspond to a uniform distribution. We introduced 12 invading groundsel plants in the center and then counted the number and position of all seedlings in two subsequent generations-hoping to determine whether the success of invasion depends on the spatial heterogeneity of the gaps. One can assess a plant’s success of invasion in two different ways. The number of individuals that get established provides one index, and the distance between a parent and its offspring-the rate of spread-provides another. In our experiments, larger gaps increased the number of established groundsel seedlings, even though we controlled for overall gap area. In addition, the invading groundsel spread faster with large gaps. For example, large gaps produced nearly three times more distance between a parent and its offspring, as compared with small gaps. Moreover, the invading groundsel produced more established seedlings with patchy gaps than uniform ones, regardless of the size of the gaps. However, offspring traveled farther when the gaps were positioned uniformly, regardless of the size of the gaps. We wondered if the way that a plant’s “shower” of seeds would fall on such gaps would lead to similar results. One can imagine that a plant produces a seed shadow, which depicts the proportion of seeds that fall relative to the distance from the plant. In our transect experiment, some seeds would fall in gaps and germinate, and others would fall in vegetation and not germinate. By knowing a plant’s seed shadow and a transect’s arrangement of gaps, one can predict the expected distance between parents and seeds that land in gaps. A simple mathematical model of this scenario produced results that resembled what we found in our experiments. In other words, how the seeds disperse and the strong competitive dominance of established grass over seedlings explains what we observed. These results point to an interesting trade-off: An invading plant can progress faster in a field that contains a uniform distribution of gaps, but fewer seeds land successfully in uniform gaps. This trade off affects models of the persistence of competitively inferior species in patchy environments. In the past, such models suggested that a competitively inferior species can persist in a community by dispersing more effectively than its superior competitor, but models of that phenomenon ignore the spatial positioning of gaps. Our results, however, indicate that a competitively inferior species faces a more difficult challenge, because of the negative relationship between rates of dispersal and the probability that seeds land in gaps and establish successfully Dispersing seeds that travel a far distance, on average, require uniformly positioned gaps; but patchy gaps lead to more established seedlings. In other words, a plant can either widely disperse its offspring or produce lots of them, but it probably cannot do both. Future research should address how competitively inferior species persist in realistic, spatially heterogeneous environments. My work with two common weedsgroundsel and bluegrass-shows that competition between these plants depends on many factors. Competition between generations-later groundsel seeds battling established bluegrass-is the primary factor that governs the dynamics of this system. Nevertheless, groundsel’s genotype determines largely when a seedling will emerge-a crucial factor in competitive success-and that suggests that contemporary plants must compete, as well. Moreover, the result of competition between groundsel and bluegrass also depends on the structure of the local environment, including the size and arrangement of gaps. In the future, ecologists hope to develop models and experimental systems that simultaneously examine how these factors contribute to plant competition. |
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