Winter low temperature disturbance in the southern subtropics of China promotes the competitiveness of an invasive plant

Winter low temperature disturbance in the southern subtropics has important effects on the weed community structure, but the role of uniquely low temperatures in biological invasions is unclear. Here, we examined the competitive effects of an invasive plant, Bidens pilosa L., and its native congener, Bidens biternata (Lour.) Merr. et Sherff, during high and low temperature seasons to determine whether low temperatures promote the competitiveness of B. pilosa in the southern subtropics of China. The growth and physiological responses of the two Bidens species to low (10/5 °C) and optimum (30/25 °C) temperatures were examined to determine how the invasive B. pilosa responds to low temperature stress. Our results showed that the competitive balance index values of B. pilosa in low temperature seasons were significantly higher than those in high temperature seasons, which implied that low temperatures may be more beneficial to the competitiveness of B. pilosa than high temperatures in the southern subtropics. The smaller decline in the relative growth rate and the photosynthetic ability of B. pilosa compared with B. biternata under low temperature stress indicated that the former was less negatively affected by low temperature than the latter. A higher DPPH^· (1.1-diphenyl-2-picrylhy-drazyl) scavenging rate and greater heat-stable protein content in B. pilosa under low temperatures might help the invasive plant to maintain more effective physiological functions and thus a higher growth rate. Overall, the uniquely low temperature in the southern subtropics of China is expected to promote the invasiveness of the exotic B. pilosa.

     

涝渍胁迫对入侵植物白花鬼针草及其本地近缘种金盏银盘生长和生理特性的影响

[目的]揭示涝渍胁迫下白花鬼针草与其本地近缘种金盏银盘的生长和生理特性差异。[方法]研究对照和涝渍胁迫下白花鬼针草和金盏银盘株高、新生不定根数目、净光合速率（P_n）、总生物量、相对生长速率、叶面积等生长和生理指标的变化。[结果]涝渍胁迫后入侵植物白花鬼针草比本地种金盏银盘保持了更高的株高、新生不定根数目和P_n（P<0.01）,更高的总生物量、相对生长速率、根生物量、茎生物量、叶生物量、叶面积（P<0.05）。涝渍胁迫10 d内,白花鬼针草根系活力和光系统II最大光量子产量显著高于金盏银盘（P<0.05）,涝渍胁迫20 d后2物种的2个参数趋于一致,这些结果表明涝渍对白花鬼针草生长的负面影响较小。涝渍胁迫前期更快的不定根生长速度和更高的根系脱氢酶活力可缓解涝渍对白花鬼针草的胁迫,减少了涝渍对光合系统的破坏,使其保持更高的P_n,从而有助于白花鬼针草保持更高的生长速率,是白花鬼针草耐受涝渍的生理机制。[结论]与本地近缘种金盏银盘相比,短期涝渍可使入侵植物白花鬼针草形成生长优势,华南地区雨季短期集中降水所形成的土壤涝渍有利于白花鬼针草的入侵。全球气候变化造成的降水不平衡性可能会加剧白花鬼针草在华南地区的入侵。     

不同光照和水分下三叶鬼针草与本地种金盏银盘生长特征的比较研究

为探讨三叶鬼针草(Bidens pilosa)成功入侵机制,利用盆栽试验对不同光照和水分条件下三叶鬼针草与本地种金盏银盘(B.biternata)的生长特征进行了比较研究。结果表明,两物种对光和水的变化均具有较大可塑性,但与金盏银盘相比,在有利环境下三叶鬼针草具有较大的株高、叶面积、生物量;在不利环境下,三叶鬼针草具有较大的比叶面积和叶面积比;除叶面积比和叶生物量比外,三叶鬼针草各生理参数的可塑性指数均高于本地种。这说明三叶鬼针草具有较强的表型可塑性和入侵适应性,这些特性促进了其成功入侵。     

Growth and leaf physiology of monkeyflowers with different altitude ranges

Every species is limited both geographically and ecologically to a subset of available habitats, yet for many species the causes of distribution limits are unknown. Temperature is thought to be one of the primary determinants of species distributions along latitudinal and altitudinal gradients. This study examined leaf physiology and plant performance under contrasting temperature regimes of sister species of monkeyflower, Mimulus cardinalis and Mimulus lewisii (Phrymaceae), that differ in altitude distribution to test the hypothesis that temperature is the primary determinant of differences in fitness versus altitude. Each species attained greatest aboveground biomass, net photosynthetic rate, and effective quantum yield of photosystem II when grown under temperatures characteristic of the altitudinal range center. Although both species exhibited greater stem length, stomatal conductance, and intercellular CO₂ concentration in hot than in cold temperatures, these traits showed much greater reductions under cold temperature for M. cardinalis than for M. lewisii. Survival of M. lewisii was also sensitive to temperature, showing a striking decrease in hot temperatures. Within each temperature regime, the species native to that temperature displayed greatest growth and leaf physiological capacity. Populations from the altitude range center and range margin of each species were used to examine population differentiation, but central and marginal populations did not differ in most growth or leaf physiological responses to temperature. This study provides evidence that M. cardinalis and M. lewisii differ in survival, growth, and leaf physiology under temperature regimes characterizing their contrasting low and high altitude range centers, and suggests that the species’ altitude range limits may arise, in part, due to metabolic limitations on growth that ultimately decrease survival and limit reproduction.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

养分增加提高大狼耙草入侵种群的生长和竞争能力

为了探讨大狼耙草的入侵风险,该文通过同质种植园实验,研究了不同养分水平下大狼耙草河北、江苏、江西和广西4个入侵种群在单种和各种群与近缘本地植物金盏银盘混种时的生长和竞争响应。结果表明:(1)单种时4个种群的株高、分枝数和总生物量在高养分下显著高于低养分下,繁殖比在低养分下显著高于高养分下(江苏种群除外); 混种时4个种群各生长参数的竞争响应在高养分下小于低养分下的。(2)各养分下,广西和江西种群的株高和总生物量显著高于河北种群,广西种群的分枝数最多[低、中和高养分下分别为(12±0.86)、(16.83±0.95)和(21.83±1.14)]; 河北种群的繁殖比在低养分 [(47.33±3.29)%]和高养分 [(25.74±2.82)%]下最高,且显著高于同养分下的广西种群 [低养分为(30.92±1.78)%和高养分为(19.77±1.22)%]。中养分下,河北种群总生物量的竞争响应(-0.51±0.04)显著大于广西种群(-0.35±0.06),繁殖生物量的竞争响应(-0.46±0.03)也显著大于广西种群(-0.28±0.07)。综上表明,高养分提高大狼耙草的生长和竞争能力,生长和竞争能力在种群间有差异,养分增加和入侵种群间基因流可能会潜在地提高大狼耙草的入侵风险,该研究结果有助于预测入侵植物的入侵风险。     

Elevated CO<Subscript>2</Subscript> increases energy-use efficiency of invasive <Emphasis Type="Italic">Wedelia trilobata</Emphasis> over its indigenous congener

Increasing atmospheric CO₂ concentration is regarded as an important factor facilitating plants invasions by stimulating invasive species growth. However, the physiological mechanisms by which invasive plants increase at the expense of existing native plants are poorly understood. Plant growth is always related to energy-use process including energy assimilation and expenditure, and thus examination of energetic properties could provide mechanistic insight into growth responses to increased CO₂. The aims of this study were to examine the effect of rising CO₂ on the growth and energetic properties of alien invasive species (Wedelia trilobata (L.) Hitchc.) and its native congener (Wedelia chinensis (Osbeck.) Merr.) in South China, and to determine if the specific energetic properties of invasive species at elevated CO₂ favoring its growth. Elevated CO₂ stimulated a greater increase in biomass production for invasive W. trilobata (58.9%) than for its indigenous congener (48.1%). Meanwhile, elevated CO₂ altered the energetic properties differently upon species. For invasive W. trilobata, elevated CO₂ significantly increased total energetic gain via photosynthetic activity (A _total), but decreased energetic cost of biomass construction (CC), and thus enhanced photosynthetic energy-use efficiency (PEUE) by 85.3%. In contrast, the indigenous W. chinensis showed a slight increase in PEUE by 43.8%. Additionally, W. trilobata individuals grown in elevated CO₂ increased energy allocation towards stems. Statistic analysis revealed significant associations between growth characteristics (relative growth rate and biomass) and energetic properties (CC and PEUE), suggesting the greater growth stimulation in invasive species could be partly explained by its specific energetic properties in elevated CO₂ concentration. The invasive species showed a greater increase in energy-use efficiency under elevated CO₂, which consequently facilitated its growth. It might be a physiological mechanism promoting success of invasion with ongoing increase in atmospheric CO₂ concentration.     

Tolerance to drought and flooding events provides a competitive advantage for an invasive over a native plant species

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Carbon and water relations of juvenile Quercus species in tall‐grass prairie

Abstract. In ecosystems where environments are extreme, such as deserts, adult plant species may facilitate the establishment and growth of seedlings and juveniles. Because high temperatures and evaporative demand characterize tall‐grass prairies of the central United States (relative to forests), we predicted that the grassland‐forest ecotone, by minimizing temperature extremes and moderating water stress, may function to facilitate the expansion of Quercus species into undisturbed tall‐grass prairie. We assessed the carbon and water relations of juvenile Quercus macrocarpa and Q. muhlenbergii, the dominant tree species in gallery forests of northeast Kansas, in ecotone and prairie sites. To evaluate the potentially competitive effects of neighboring herbaceous biomass on these oaks, juveniles (< 0.5 m tall) of both species also were subjected to either: (1) removal of surrounding above‐ground herbaceous biomass, or (2) control (prairie community intact) treatments. Herbaceous biomass removal had no significant effect on gas exchange or water relations in these oak species in either the prairie or the ecotone environment. Although the ecotone did alleviate some environmental extremes, photosynthetic rates and stomatal conductance were ca. 20 % higher (p < 0.05) in both oaks in prairie sites vs. the ecotone. Moreover, although leaf temperatures on average were higher in oaks in the prairie, high leaf temperatures in the ecotone had a greater negative effect on photosynthesis. These data suggest that the grassland‐forest ecotone did not facilitate the growth of Quercus juveniles expanding into this grassland. Moreover, the carbon and water relations of juvenile oaks in the prairie appeared to be unaffected by the presence of the dominant C₄ grasses.     

Seasonal and diurnal patterns of photosynthetic gas exchange for Lonicera sempervirens and L. Japonica (Caprifoliaceae)

Photosynthesis, stomatal conductance, and water use efficiency were compared between Lonicera japonica (Japanese honeysuckle), an invasive species in the southeastern United States, and its native congener, Lonicera sempervirens (coral honeysuckle), to determine the role of seasonal patterns of water loss and carbon gain in the invasive ability of the exotic. Diurnal measurements were taken monthly for 1 year under a closed forest canopy and in an open field. There were few significant differences in photosynthetic rates between the two species in either environment. However, at both sites, Lonicera japonica retained its old leaves over winter while old leaves of L. sempervirens senesced. Also, new leaves of L. japonica had significantly higher photosynthetic rates than the emerging leaves of L. sempervirens (6.2 vs. 4.4 μmol m ²sec^-1 under the canopy; 4.4 vs. 3.0 μmol m^-2 sec^-1 in the open). Although differences in conductance and water use efficiency between species were seldom significant, L. japonica tended to have higher maximum values than L. sempervirens. Retention of old leaves by L. japonica during new leaf formation (January–March) as well as higher photosynthetic rates in new leaves contribute to greater annual carbon gain and help explain the invasive ability of Japanese honeysuckle.     

Photosynthesis and water‐use efficiency: A comparison between invasive (exotic) and non‐invasive (native) species

Abstract Invasive species have been hypothesized to out‐compete natives though either a Jack‐of‐all‐trades strategy, where they are able to utilize resources effectively in unfavourable environments, a master‐of‐some, where resource utilization is greater than its competitors in favourable environments, or a combination of the two (Jack‐and‐master). We examined the invasive strategy of Berberis darwinii in New Zealand compared with four co‐occurring native species by examining germination, seedling survival, photosynthetic characteristics and water‐use efficiency of adult plants, in sun and shade environments. Berberis darwinii seeds germinated more in shady sites than the other natives, but survival was low. In contrast, while germination of B. darwinii was the same as the native species in sunny sites, seedling survival after 18 months was nearly twice that of the all native species. The maximum photosynthetic rate of B. darwinii was nearly double that of all native species in the sun, but was similar among all species in the shade. Other photosynthetic traits (quantum yield and stomatal conductance) did not generally differ between B. darwinii and the native species, regardless of light environment. Berberis darwinii had more positive values of δ¹³C than the four native species, suggesting that it gains more carbon per unit water transpired than the competing native species. These results suggest that the invasion success of B. darwinii may be partially explained by combination of a Jack‐of‐all‐trades scenario of widespread germination with a master‐of‐some scenario through its ability to photosynthesize at higher rates in the sun and, hence, gain a rapid height and biomass advantage over native species in favourable environments.     

PHOTOSYNTHETIC SENSITIVITY TO TEMPERATURE IN POPULATIONS OF TWO C4 BOUTELOUA (POACEAE) SPECIES NATIVE TO DIFFERENT ALTITUDES

The relationships between photosynthesis, flowering, and growth temperatures were examined experimentally in four populations of the C₄ grass genus Bouteloua. Field-collected plants were grown under two temperature regimes, cool (20 C day/6 C night) and warm (30/16), representative of the extreme populations. Populations collected from the warm climates had significantly lower photosynthetic capacity when grown in the cool chamber relative to the warm chamber, while photosynthetic capacity in the cool climate populations did not differ between the growth conditions. Additionally, exposure to a 2-day cold temperature treatment (10/-2), representative of late-season frosts in high altitude sites, resulted in further reductions in photosynthesis in the warm climate plants, but not in the cool climate plants. This effect was greater for plants grown in the cool growth chamber. Flowering was reduced by 70% in the warm climate plants grown in the cool chamber, and was correlated with photosynthetic inhibition following the short-term cold temperature treatment. These results indicate that genetic differentiation for photosynthetic temperature sensitivity has occurred in the cool climate populations, and that long-term exposure to cool temperatures coupled with short-term relatively extreme low temperatures results in greater photosynthetic inhibition in nontolerant populations.     

Measures of ecological association

Summary The relationships of photosynthetic characteristics to the competitive interactions of a C₃ plant, Chenopodium album, and a C₄ plant, Amaranthis retroflexus, were investigated in different temperature and water supply regimes. Both species had similar photosynthetic rates at 25°C, but at higher temperatures, Amaranthus had substantially greater rates than Chenopodium. Conversely, at lower temperatures, Chenopodium had an advantage. The competitive abilities in mixtures exhibited a close parallel to the photosynthetic performances with Amaranthus having an advantage at high temperatures and Chenopodium an advantage at low temperatures. These competitive outcomes were determined primarily by differences in relative growth rates prior to canopy closure. In the respective, temperature regimes, the species having the highest photosynthetic rate, which resulted an more rapid growth, overtopped and shaded the other species at the time of canopy closure. These results demonstrate that differences in photosynthetic temperature response between C₄ and C₃ plants can be an important determinant in competitive interactions, but at least in this case, the influence is primarily through, events prior to the actual initiation of competition.In contrast to temperature, growth of the plants under limited water supply had no influence on the competitive interactions. Thus, the presence of the C₄ pathway alone was not sufficient to yield a competitive advantage over the C₃ species under water limited conditions.     

The influence of water stress on the photosynthetic performance and stomatal behaviour of tree seedlings subjected to variation in temperature and irradiance

Summary Seedlings of Betula pendula Roth. and Gmelina arborea L. were subjected to variation in temperature and irradiance. The influence of a mild water-stressing treatment on the photosynthetic performance and stomatal behaviour of these plants was assessed. For both species, the shape of the relationships between irradiance and photosynthesis and temperature and photosynthesis resembled those reported for other species. The effect of water stress was to reduce the rate of photosynthesis, particularly at high temperatures. This was largely a function of a reduction in mesophyll conductance under these conditions. The optimum temperature for stomatal opening was significantly lower than the optimum temperature for photosynthesis, which was in turn lowered by the water stress treatment. The stomata of birch seedlings showed maximum opening at an intermediate temperature while the stomata of Gmelina generally exhibited a closing movement when leaf temperatures increased from 15° C. Mesophyll conductances of both species increased with increasing temperature.The physiological basis for the variation in photosynthetic performance and stomatal behaviour and the ecological significance of this variation are discussed.     

陕北水蚀风蚀交错带柠条适应不同生境的形态和生理可塑性

以陕北水蚀风蚀交错带4个不同坡位与土壤质地生境[沟底+坝淤绵沙土(A)、梁坡+红黄土(B)、坡顶+绵沙土(C)、坡顶+风沙土盖绵沙土(D)]中生长的柠条为研究对象,研究了其生长、光合和水力性状的可塑性变化,以揭示柠条对不同生境的适应机制。结果表明:(1)生境D和生境A样地1~3m和3m土层的平均含水量明显高于生境B和C;(2)与生境B和C相比,生境A和D中生长的柠条冠幅、株高和新枝长显著增加,叶厚度减小,正午叶水势、净光合速率、气孔导度和蒸腾速率亦显著增加,但不同生境中柠条枝比导水率、Huber值和栓塞程度无显著差异;(3)柠条各测定指标中,新枝长、净光合速率、气孔导度和蒸腾速率表现出较大的可塑性。研究认为,柠条可能主要通过新枝长或光合生理特征的改变来适应不同的水分生境。     

Growth, biomass allocation and photosynthesis of invasive and native Hawaiian rainforest species

Growth, biomass allocation, and photosynthetic characteristics of seedlings of five invasive non-indigenous and four native species grown under different light regimes were studied to help explain the success of invasive species in Hawaiian rainforests. Plants were grown under three greenhouse light levels representative of those found in the center and edge of gaps and in the understory of Hawaiian rainforests, and under an additional treatment with unaltered shade. Relative growth rates (RGRs) of invasive species grown in sun and partial shade were significantly higher than those for native species, averaging 0.25 and 0.17 g g⁻¹ week⁻¹, respectively, while native species averaged only 0.09 and 0.06 g g⁻¹ week⁻¹, respectively. The RGR of invasive species under the shade treatment was 40% higher than that of native species. Leaf area ratios (LARs) of sun and partial-shade-grown invasive and native species were similar but the LAR of invasive species in the shade was, on average, 20% higher than that of native species. There were no differences between invasive and native species in biomass allocation to shoots and roots, or in leaf mass per area across light environments. Light-saturated photosynthetic rates (Pmax) were higher for invasive species than for native species in all light treatments. Pmax of invasive species grown in the sun treatment, for example, ranged from 5.5 to 11.9 μmol m⁻² s⁻¹ as compared with 3.0−4.5 μmol m⁻² s⁻¹ for native species grown under similar light conditions. The slope of the linear relationship between Pmax and dark respiration was steeper for invasive than for native species, indicating that invasive species assimilate more CO₂ at a lower respiratory cost than native species. These results suggest that the invasive species may have higher growth rates than the native species as a consequence of higher photosynthetic capacities under sun and partial shade, lower dark respiration under all light treatments, and higher LARs when growing under shade conditions. Overall, invasive species appear to be better suited than native species to capturing and utilizing light resources, particularly in high-light environments such as those characterized by relatively high levels of disturbance. Received: 30 December 1997 / Accepted: 1 September 1998     

Warmer and less variable temperatures favour an accelerated plant phenology of two invasive weeds across sub‐Antarctic Macquarie Island

The great plasticity and diverse reproductive strategies of invasive alien plants are widely assumed to contribute to invasion success, even in extreme areas, often displacing native species. In this context, climate change creates new opportunities for biological invasions. Environmental variability and global warming are two of the climatic processes that may promote invasiveness, since alien species modulate their phenology to succeed under these circumstances. We monitored the phenological development (phenological stage advancement) of the two main invasive alien species: Poa annua L. and Cerastium fontanum Baumg. in the sub‐Antarctic Macquarie Island during the austral summer period along an altitudinal gradient. We found that higher temperatures lead to increased plant height and accelerated phenological development than lower temperatures in P. annua but found no direct evidence of the latter in C. fontanum. However, increased temperature variability negatively affected the phenological development of both species. Interestingly, despite their different reproductive strategy (rapid and impromptu in P. annua, and more synchronic and gradual in C. fontanum), both species prolifically succeeded in producing seeds at all sites showing the great acclimation of these two alien species even in limiting conditions. Since both alien species in Macquarie Island showed larger size and faster phenology at lower altitudes (i.e. milder conditions), this would indicate a great influence of ameliorating abiotic extremes on alien plant invasive capabilities at environmental extremes. Thus, our results warn of the increasing capabilities under climatic warming scenarios for alien plants to reproduce even at such remote ranges. This highlights the need to reinforce calls for special attention to prevent the spread of these kinds of species to other similar sub‐polar areas, where intensive post‐introduction management may be difficult or expensive.     

遮荫处理对秋枫幼苗生理生态特性的影响

为了解遮荫对秋枫(Bischofia javanica)幼苗生长的影响,对夏季1年生秋枫幼苗在4种遮荫(透光率分别为自然光的100%、41.3%、14.6%和3.6%)处理150 d后的生理指标变化进行了研究。结果表明,1年生秋枫幼苗的光补偿点(LCP)、光饱和点(LSP)和暗呼吸速率(Rd)随着遮荫程度加重而减小,表观量子效率(AQY)则增大,净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和胞间CO_2浓度(Ci)的日变化表明秋枫有明显的"午休"现象,遮荫引起光合速率降低的主要原因是非气孔限制,而全光照条件下则是气孔限制。叶绿素、可溶性糖含量随遮荫程度的增大呈先升后降的变化趋势;相对电导率、丙二醛(MDA)含量、过氧化氢酶(CAT)活性则呈先降后升的趋势,以41.3%透光率的最小;可溶性蛋白质含量、超氧化物歧化酶(SOD)活性随遮荫程度的增大而降低。因此,秋枫幼苗主要通过提高Pn、抗氧化酶活性(SOD和CAT)、叶绿素含量,降低LSP、LCP和Rd,增大AQY来提高弱光利用能力;适度遮荫(41.3%NS)更有利于1年生秋枫幼苗在夏季的生长。     

Will climate warming exceed lethal photosynthetic temperature thresholds of lichens in a southern African arid region?

Predicted elevated temperatures and a shift from a winter to summer rainfall pattern associated with global warming could result in the exposure of hydrated lichens during summer to more numerous temperature extremes that exceed their thermal thresholds. This hypothesis was tested by measuring lethal temperature thresholds under laboratory and natural conditions for four epilithic lichen species (Xanthoparmelia austro‐africana, X. hyporhytida, Xanthoparmelia sp., Xanthomaculina hottentotta) occurring on quartz gravel substrates at a hot arid inland site two epigeous lichen species (Teloschistes capensis, Ramalina sp.) occurring on gypsum‐rich topsoil at a warm humid coastal site. Extrapolated lethal temperatures for photosynthetic quantum yield under laboratory conditions were up to 4°C higher for lichens from a dry inland site than those from a humid coastal site. Lethal temperatures extrapolated for photosynthetic quantum yield at a saturating photosynthetic photon flux density of ≥11,000 μmol photons m^?2 s^?1 under natural conditions were up to 6°C higher for lichens from the dry inland site than the more humid coastal site. It is concluded that only under atypical conditions of lichen exposure in a hydrated state to temperature extremes at high midday solar irradiances during summer could lethal photosynthetic thresholds in sensitive lichen species be potentially exceeded, but whether the increased frequency of such conditions with climate warming would lead to increased likelihood of lichen mortality is debatable.     

Contrasting hypoxia tolerance and adaptation in Malus species is linked to differences in stomatal behavior and photosynthesis

We examined the potential differences in tolerance to hypoxia by two species of apple rootstocks. Stomatal behavior and photosynthesis were compared between Malus sieversii and Malus hupehensis. Plants were hydroponically grown for 15 days in normoxic or hypoxic nutrient solutions. Those of M. sieversii showed much greater sensitivity, with exposure to hypoxia resulting in higher leaf concentrations of abscisic acid (ABA) that prompted stomatal closure. Compared with the control plants of that species, stomatal density was greater in both new and mature leaves under stress conditions. In contrast, stomatal density was significantly decreased in leaves from M. hupehensis, while stomatal length was unaffected. Under stress, the net photosynthetic rate, stomatal conductance and chlorophyll contents were markedly reduced in M. sieversii. The relatively hypoxia‐tolerant genotype M. hupehensis, however, showed only minor changes in net photosynthesis or chlorophyll content, and only a slight decrease in stomatal conductance due to such treatment. Therefore, we conclude that the more tolerant M. hupehensis utilizes a better protective mechanism for retaining higher photosynthetic capacity than does the hypoxia‐sensitive M. sieversii. Moreover, this contrast in tolerance and adaptation to stress is linked to differences in their stomatal behavior, photosynthetic capacity and possibly their patterns of native distribution.     

Innate and evolutionarily increased advantages of invasive <Emphasis Type="Italic">Eupatorium adenophorum</Emphasis> over native <Emphasis Type="Italic">E. japonicum</Emphasis> under ambient and doubled atmospheric CO<Subscript>2</Subscript> concentrations

Both innate and evolutionarily increased ecophysiological advantages can contribute to vigorous growth, and eventually to invasiveness of alien plants. Little effort has been made to explore the roles of innate factors of alien plants in invasiveness and the effects of CO₂ enrichment on alien plant invasions. To address these problems, we compared invasive Eupatorium adenophorum, its native conspecific, and a native congener (E. japonicum) under ambient and doubled atmospheric CO₂ concentrations. Native E. adenophorum from Mexico grew slower than invasive E. adenophorum but faster than native E. japonicum under both CO₂ concentrations. The faster growth rate of invasive E. adenophorum was associated with higher photosynthetic capacity and leaf area ratio. For invasive E. adenophorum, the higher photosynthetic capacity was associated with higher nitrogen (N) allocation to photosynthesis, which was related to lower leaf mass per area; the higher leaf area ratio was due to lower leaf mass per area and higher leaf mass fraction. Tradeoff between N allocations to photosynthesis versus defenses was found. CO₂ enrichment significantly increased relative growth rate and biomass accumulation by increasing actual photosynthetic rate for all studied materials. However, the relative increase in growth was not significantly different among them. CO₂ enrichment did not influence N allocation to photosynthesis, but increased N allocation to cell walls. The reduced leaf N content decreased N content in photosynthesis, explaining the down-regulation of photosynthetic capacity under prolonged elevated CO₂ concentration. Our results indicate that both innate and evolutionary advantages in growth and related ecophysiological traits contribute to invasiveness of invasive E. adenophorum, and CO₂ enrichment may not aggravate E. adenophroum’s invasion in the future.