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Invasive species can change selective pressures on native plants by altering biotic and abiotic conditions in invaded habitats. Although invasions can lead to native species extirpation, they may also induce rapid evolutionary changes in remnant native plants. We investigated whether adult plants of five native perennial grasses exhibited trait shifts consistent with evolution in response to invasion by the introduced annual grass Bromus tectorum L. (cheatgrass), and asked how much variation there was among species and populations in the ability to grow successfully with the invader. Three hundred and twenty adult plants were collected from invaded and uninvaded communities from four locations near Reno, Nevada, USA. Each plant was divided in two and transplanted into the greenhouse. One clone was grown with B. tectorum while the other was grown alone, and we measured tolerance (ability to maintain size) and the ability to reduce size of B. tectorum for each plant. Plants from invaded populations consistently had earlier phenology than those from uninvaded populations, and in two out of four sites, invaded populations were more tolerant of B. tectorum competition than uninvaded populations. Poa secunda and one population of E. multisetus had the strongest suppressive effect on B. tectorum, and these two species were the only ones that flowered in competition with B. tectorum. Our study indicates that response to B. tectorum is a function of both location and species identity, with some, but not all, populations of native grasses showing trait shifts consistent with evolution in response to B. tectorum invasion within the Great Basin.

The life-long addition of new neurons has been documented in many regions of the vertebrate and invertebrate brain, including the hippocampus of mammals (Altman and Das, 1965; Eriksson et al., 1998; Jacobs et al., 2000), song control nuclei of birds (Alvarez-Buylla et al., 1990), and olfactory pathway of rodents (Lois and Alvarez-Buylla, 1994), insects (Cayre et al., 1996) and crustaceans (Harzsch and Dawirs, 1996; Sandeman et al., 1998; Harzsch et al., 1999; Schmidt, 2001). The possibility of persistent neurogenesis in the neocortex of primates is also being widely discussed (Gould et al., 1999; Kornack and Rakic, 2001). In these systems, an effort is underway to understand the regulatory mechanisms that control the timing and rate of neurogenesis. Hormonal cycles (Rasika et al., 1994; Harrison et al., 2001), serotonin (Gould, 1999; Brezun and Daszuta, 2000; Beltz et al., 2001), physical activity (Van Praag et al., 1999) and living conditions (Kemperman and Gage, 1999; Sandeman and Sandeman, 2000) influence the rate of neuronal proliferation and survival in a variety of organisms, suggesting that mechanisms controlling life-long neurogenesis are conserved across a range of vertebrate and invertebrate species. The present article extends these findings by demonstrating circadian control of neurogenesis. Data show a diurnal rhythm of neurogenesis among the olfactory projection neurons in the crustacean brain, with peak proliferation during the hours surrounding dusk, the most active period for lobsters. These data raise the possibility that light-controlled rhythms are a primary regulator of neuronal proliferation, and that previously-demonstrated hormonal and activity-driven influences over neurogenesis may be secondary events in a complex circadian control pathway.

Introduced grasses have displaced Hawaiian Pili grass (Heteropogon contortus) in most dry, leeward habitats of the Hawaiian Islands. The purpose of this study was to assess the feasibility of restoring an indigenous Heteropogon grassland at the Puukohola Heiau National Historic Site, where introduced Buffel grass (Cenchrus ciliaris) is now dominant. Heteropogon seeds (50 seedlings/m 2 ) were added to replicate plots within a Cenchrus grassland. Some plots were subjected to one-time herbicide or hand-pulling treatments to remove established Cenchrus. Because Hawaiians historically used fire to promote Heteropogon grasslands, the plots were burned biennially. Plots were also subjected to two levels of water supplementation. Heteropogon establishment was monitored over 2 and 4 years in the higher-and lower-water plots, respectively. In treatments containing established Cenchrus, Heteropogon establishment was consistently poor (<10% cover). But in the burned plots where established Cenchrus had been removed, as many as 31 Heteropogon seedlings per square meter were recorded, and Heteropogon became the dominant cover, averaging 34% absolute cover (81% relative cover) after 4 years in the lower-water plot and 34% absolute cover (60% relative cover) after 2 years in the higherwater plot. Few Cenchrus grass seedlings survived, possibly due to insufficient water. Water supplementation promoted growth of other alien grasses from the seed bank (Digitaria insularis and Eragrostis spp.); however, these grasses quickly declined after supplemental watering was terminated. Although initial suppression of Cenchrus was required, Heteropogon expanded quickly when seeds and fire were reintroduced, demonstrating that a Heteropogondominated grassland can be reestablished in 2-4 years.

New neurons are incorporated into the adult brains of a variety of organisms, from humans and higher vertebrates, to non-vertebrates such as crustaceans. In virtually all of these systems serotonergic pathways appear to provide important regulatory influences over the machinery producing the new neurons. We have developed an in vitro preparation where adult neurogenesis can be maintained under highly controlled conditions, and are using this to test the influence of hormones on the production of neurons in the crustacean (Homarus americanus) brain. Serotonin levels have been manipulated in this in vitro preparation, and the resulting effects on the rate of neurogenesis have been documented. In addition we have compared in vitro influences of serotonin with results acquired from in vivo exposure of whole animals to serotonin. These experiments suggest that there are multiple mechanisms and pathways by which serotonin may regulate neurogenesis in the crustacean brain: (1) serotonin is effective in regulating neurogenesis at levels as low as 10−10M, suggesting that circulating serotonin may have hormonal influences on neuronal precursor cells residing in the proliferation zones; (2) contrasting effects of serotonin on neurogenesis (up- vs. down-regulation) at high concentrations (10−4M), dependent upon whether eyestalk tissue is present or absent, indicate that serotonin elicits the release of substances from the sinus glands that are capable of suppressing neurogenesis; (3) previously demonstrated (Beltz et al., 2001) serotonergic fibers from the dorsal giant neuron project directly into the proliferation zone in Cluster 10, suggest synaptic or local influences on neurogenesis in the proliferation zones where the final cell divisions and neuronal differentiation occur. Serotonin therefore regulates neurogenesis by multiple pathways, and the specific mode of influence is concentration-dependent.

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Summary While much research has documented the impact of invaders on native communities and ecosystem services, there has been less work quantifying how invasion affects the genetic composition of native populations. That is, when invaders dominate a community, can they shift selection regimes and impact the evolutionary trajectory of native populations? The invasion of the annual grass Bromus tectorum in the Intermountain West provides an opportunity to quantify the effects of invasion on natural selection in wild populations. The shift from a perennial‐dominated native community to one dominated by a highly competitive annual species alters the timing and intensity of competitive pressure, which has the potential to strongly shift selection regimes for native plants. To quantify traits under selection in contrasting environments, we planted seeds of two native perennial grasses, Elymus multisetus and Poa secunda, into three invaded, invaded but weeded and relatively uninvaded sagebrush systems. We quantified phenotypic traits of seedlings from separate maternal plants, describing differences in phenotypes among individuals. We then asked which traits were associated with survival and plant size in adjacent invaded and uninvaded sagebrush systems, following individual seed performance for 3 years. We found evidence for divergent selection between invaded and uninvaded sagebrush systems, with contrasting phenotypic traits associated with greater survival or plant size in these different growing conditions at all three field sites. For example, at one field site, P. secunda families with higher root tip production were more likely to survive in invaded and weeded environments, but this was not the case in uninvaded environments. Similarly, for E. multisetus, root mass fraction, seed mass and allocation to coarse or fine roots affected survival and plant size, again with contrasting relationships across invaded, weeded or uninvaded environments. Synthesis. Impacts of invasive species extend beyond ecosystem and community composition changes and can affect the evolutionary trajectory of native populations. By quantifying natural selection in invaded landscapes, we identified phenotypic traits that are potentially adaptive in these invaded systems. Importantly, these traits differed from traits associated with success in uninvaded communities. This insight into adaptive, contemporary evolution in native species can guide restoration and conservation efforts.

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