Phenotypic plasticity and genetic variation in leaf traits of Yushania niitakayamensis (Bambusoideae; Poaceae) in contrasting light environments.

Yushania niitakayamensis (Bambusoideae; Poaceae), a perennial grass distributed from mid to high elevations in Taiwan, is often found growing in exposed grassland or shaded forest understories. To answer the question how does Y. niitakayamensis cope with contrasting light availability of habitats, we compared (1) leaf ecophysiological traits between populations of Y. niitakayamensis growing in exposed and shaded habitats in four seasons, and (2) plasticity patterns of transplanted plants to two light treatments (full-sun and 70 % shading) in a phytotron. Significant differences in leaf morphological (leaf length, width, specific leaf area, stomatal density), anatomical (leaf thickness (LT), the frequency of cavity formed by collapsed fusoid cells), and biochemical (chlorophyll contents, nitrogen contents per unit area ([N]area) and stable carbon isotope ratio) features were found between populations across seasons. Common garden experiments suggested that most of the trait variations in field growing plants can be explained by the effect of growth light treatment but not by that of population. However, variations between the two populations in LT, [N]area, gas exchange parameters, and the degree of plasticity in LT and [N]area in responding to growth light regimes might have genetic basis. In comparison between transplants from different origins grown under same light regime, plants from the exposed population grown under full-sun expressed significantly higher LT, [N]area and light-saturated photosynthetic rates, whereas plants from the shaded population grown under shading had significantly higher water use efficiency. Accordingly, local specialization in populations of Y. niitakayamensis to particular environmental conditions might have arisen.

Phylogenetic analyses of Bradyrhizobium symbionts associated with invasive Crotalaria zanzibarica and its coexisting legumes in Taiwan.

In this study, the genetic diversity and identification of Bradyrhizobium symbionts of Crotalaria zanzibarica, the most widely-distributed invasive legume in Taiwan, and other sympatric legume species growing along riverbanks of Taiwan were evaluated for the first time. In total, 59 and 54 Bradyrhizobium isolates were obtained from C. zanzibarica and its coexisting legume species, respectively. Based on the multilocus sequence analysis (MLSA) of concatenated four housekeeping genes (dnaK-glnII-recA-rpoB gene sequences, 1901bp), the 113 isolates displayed 53 unique haplotypes and grouped into 21 clades. Of these clades, 11 were found to be congruent to already defined Bradyrhizobium species, while the other 10 clades were found to not be congruent to any defined species. In particular, the C. zanzibarica isolates belong to 14 MLSA clades, six of which overlapped with the isolates of coexisting legumes. According to the nodA gene sequences (555bp) obtained from the 105 isolates, these isolates were classified into three known nodA clades, III.2, III.3 and VII and were further clustered into 10 groups. Furthermore, the C. zanzibarica isolates were clustered into 8 nodA groups, five of which overlapped with the isolates from coexisting legumes. Additionally, the nodA genes of the isolates from native species were dominated by Asian origin, while those from C. zanzibarica were dominated by American origin. In conclusion, C. zanzibarica is a promiscuous host capable of recruiting diverse Bradyrhizobium symbionts, some of which are phylogenetically similar to the symbionts of coexisting legumes in Taiwan.

Phylogenetic Identification, Phenotypic Variations, and Symbiotic Characteristics of the Peculiar Rhizobium, Strain CzR2, Isolated from Crotalaria zanzibarica in Taiwan.

Crotalaria zanzibarica is an exotic and widely distributed leguminous plant in Taiwan. The relationship between C. zanzibarica and its rhizobial symbionts has been suggested to contribute to its successful invasion. A rhizobial strain (designed as CzR2) isolated from the root nodules of C. zanzibarica and cultivated in standard YEM medium displayed pleomorphism, with cells ranging between 2 and 10 µm in length and some branching. In the present study, we identified this rhizobial strain, investigated the causes of pleomorphism, and examined the nodules formed. The results of a multilocus sequence analysis of the atpD, dnaK, glnII, gyrB, recA, and rpoB genes revealed that CzR2 belongs to Bradyrhizobium arachidis, a peanut symbiont recently isolated from China. Cells of the strain were uniformly rod-shaped in basal HM medium, but displayed pleomorphism in the presence of yeast extract, mannitol, or fructose. These results indicate that the morphology of CzR2 in its free-living state is affected by nutrient conditions. Several highly pleomorphic bacteroids enclosed in symbiosomes were frequently detected in FM and TEM observations of sections of the indeterminate nodules induced by CzR2; however, no infection thread was identified. Flow cytometric analyses showed that CzR2 cells in YEM medium and in the nodules of C. zanzibarica had two or more than two peaks in relative DNA contents, respectively, suggesting that the elongated cells of CzR2 in its free-living state occur due to a cell cycle-delayed process, while those in its symbiotic state are from genomic endo-reduplication.

Chromosome numbers of populations of three varieties of Bidens pilosa in Taiwan.

BACKGROUND: Hairy beggar-ticks (Bidens pilosa L.) is a common invasive plant in tropical and subtropical regions. The Flora of Taiwan listed three varieties of B. pilosa in Taiwan, var. minor, var. pilosa and var. radiata. Among the three varieties, var. radiata was the most recently, in 1970s, introduced into Taiwan. However, after its introduction into Taiwan, var. radiata has become dominant over the other two varieties and is considered a serious invasive plant in lowland of Taiwan. Our previous study showed that var. radiata is self-incompatible and the other two varieties are self-fertile. Could it be possible that different chromosome numbers contribute to the different breeding systems of these three varieties? In addition, the heterogeneities of traits of var. radiata were found higher than those of var. minor and var. pilosa. Is the phenomenon resulting from the hybridization between var. radiata with other varieties? We counted chromosome numbers of populations of these three varieties distributed in Taiwan and conducted hand pollination treatment between var. radiata (as pollen receiver) and var. minor or var. pilosa (as pollen donor) to provide answer for the aforementioned questions. RESULTS: No difference was found in chromosome numbers among populations of the same variety. Forty-eight chromosomes (2n = 48) were counted for var. radiata while 72 (2n = 72) chromosomes for var. minor and var. pilosa. Therefore, var. radiata is tetraploid and var. minor and var. pilosa are hexaploid. No successful hybridization was found between var. radiata and var. minor or between var. radiata and var. pilosa. CONCLUSIONS: This study provided the evidence that the invasive plant (B. pilosa var. radiata) has different chromosome numbers from the other two varieties and is unlikely to hybridize with the other two varieties.

Phototropic leaf movements and photosynthetic performance in an amphibious fern, Marsilea quadrifolia.

Diurnal phototropism has not been reported in ferns. In this study we found that the four leaflets of the amphibious fern Marsilea quadrifolia are capable of adjusting their leaflet angle and leaflet azimuth in response to changes in the position of the sun's direct beam, exhibiting more diaphototropic movements (orienting the plane of the lamina perpendicular to incident light) in the morning and late afternoon, and more paraphototropic movements (orienting the plane of the lamina parallel to incident light) at noon. In addition, by cutting off the leaflet lamina and covering portions of leaflets with black tape, the junction between the leaflet and petiole was found to be responsible for light reception. Among the light spectrum investigated, blue light was the most effective at inducing diaphototropism. The role of diurnal phototropism in enhancing carbon return and ameliorating photoinhibition was also evaluated. It was concluded that diurnal phototropic leaf movement represents one of the plastic responses enabling this amphibious fern to grow under terrestrial conditions.

Sensitivity to chilling temperatures and distribution differ in the mangrove species Kandelia candel and Avicennia marina.

We compared the effects of short-term (hours) and long-term (days) exposure to chilling temperatures on the photosynthetic gas exchange, leaf characteristics and chlorophyll a fluorescence of seedlings of the mangrove species Kandelia candel Druce and Avicennia marina (Forsk.) Vierh. Both species occur along the west coast of Taiwan, but K. candel occurs further north than A. marina. We hypothesized that temperature was one of the major environmental factors limiting the northern distribution of A. marina. Avicennia marina was more sensitive to chilling temperatures than K. candel. Leaves of both species showed reductions in light-saturated photosynthetic rates (Amax), stomatal conductance (gs) and quantum yield of photosystem II after a 1-h exposure to 15 degrees C, with A. marina showing significantly greater reductions in Amax and gs than K. candel. No significant differences in Amax, gs and electron transport rate (ETR) were found between leaves of K. candel grown at 15 and 30 degrees C for 10 days. However, leaves of A. marina grown for 10 days at 15 degrees C had significantly lower Amax, gs and ETR than plants grown at 30 degrees C. After 20 days at 15 degrees C, leaf mass per area of both species was increased significantly, whereas area-based chlorophyll concentrations were reduced, with significantly greater changes in A. marina than in K. candel. We concluded that sensitivity to low winter temperatures is a primary limiting factor in the distribution of A. marina along the western coast of Taiwan.

Influence of stand structure on carbon-13 of vegetation, soils, and canopy air within deciduous and evergreen forests in Utah, United States.

Carbon isotope ratios (δ13C) were studied in evergreen and deciduous forest ecosystems in semi-arid Utah (Pinus contorta, Populus tremuloides, Acer negundo and Acer grandidentatum). Measurements were taken in four to five stands of each forest ecosystem differing in overstory leaf area index (LAI) during two consecutive growing seasons. The δ13Cleaf (and carbon isotope discrimination) of understory vegetation in the evergreen stands (LAI 1.5-2.2) did not differ among canopies with increasing LAI, whereas understory in the deciduous stands (LAI 1.5-4.5) exhibited strongly decreasing δ13Cleaf values (increasing carbon isotope discrimination) with increasing LAI. The δ13C values of needles and leaves at the top of the canopy were relatively constant over the entire LAI range, indicating no change in intrinsic water-use efficiency with overstory LAI. In all canopies, δ13Cleaf decreased with decreasing height above the forest floor, primarily due to physiological changes affecting c i/c a (> 60%) and to a minor extent due to δ13C of canopy air (< 40%). This intra-canopy depletion of δ13Cleaf was lowest in the open stand (1‰) and greatest in the denser stands (4.5‰). Although overstory δ13Cleaf did not change with canopy LAI, δ13C of soil organic carbon increased with increasing LAI in Pinus contorta and Populus tremuloides ecosystems. In addition, δ13C of decomposing organic carbon became increasingly enriched over time (by 1.7-2.9‰) for all deciduous and evergreen dry temperate forests. The δ13Ccanopy of CO2 in canopy air varied temporally and spatially in all forest stands. Vertical canopy gradients of δ13Ccanopy, and [CO2]canopy were larger in the deciduous Populus tremuloides than in the evergreen Pinu contorta stands of similar LAI. In a very wet and cool year, ecosystem discrimination (Δe) was similar for both deciduous Populus tremulodies (18.0 ± 0.7‰) and evergreen Pinus contorta (18.3 ± 0.9‰) stands. Gradients of δ13Ccanopy and [CO2]canopy were larger in denser Acer spp. stands than those in the open stand. However, 13C enrichment above and photosynthetic draw-down of [CO2]canopy below tropospheric baseline values were larger in the open than in the dense stands, due to the presence of a vigorous understory vegetation. Seasonal patterns of the relationship δ13Ccanopy versus 1/[CO2]canopy were strongly influenced by precipitation and air temperature during the growing season. Estimates of Δe for Acer spp. did not show a significant effect of stand structure, and averaged 16.8 ± 0.5‰ in 1933 and 17.4 ± 0.7‰ in 1994. However, Δe varied seasonally with small fluctuations for the open stand (2‰), but more pronounced changes for the dense stand (5‰).