Inter- and intra-generic differences in growth,reproduction, and fitness of nine herbaceous annual species grown in elevated CO2 environments

In assessing the capacity of plants to adapt to rapidly changing global climate, we must elucidate the impacts of elevated carbon dioxide on reproduction, fitness and evolution. We investigated how elevated CO₂ influenced reproduction and growth of plants exhibiting a range of floral morphologies, the implications of shifts in allocation for fitness in these species, and whether related taxa would show similar patterns of response. Three herbaceous, annual species each of the genera Polygonum, Ipomoea, and Cassia were grown under 350 or 700 ppm CO₂. Vegetative growth and reproductive output were measured non-destructively throughout the full life span, and vegetative biomass was quantified for a subsample of plants in a harvest at first flowering. Viability and germination studies of seed progeny were conducted to characterize fitness precisely. Early vegetative growth was often enhanced in high-CO₂ grown plants of Polygonum and Cassia (but not Ipomoea). However, early vegetative growth was not a strong predictor of subsequent reproduction. Phenology and production of floral buds, flowers, unripe and abscised fruits differed between CO₂ treatments, and genera differed in their reproductive and fitness responses to elevated CO₂. Polygonum and Cassia species showed accelerated, enhanced reproduction, while Ipomoea species generally declined in reproductive output in elevated CO₂. Seed quality and fitness (in terms of viability and percentage germination) were not always directly correlated with quantity produced, indicating that output alone may not reliably indicate fitness or evolutionary potential. Species within genera typically responded more consistently to CO₂ than unrelated species. Cluster analyses were performed separately on suites of vegetative and reproductive characters. Some species assorted within genera when these reproductive responses were considered, but vegetative responses did not reflect taxonomic affinity in these plants. Congeners may respond similarly in terms of reproductive output under global change, but fitness and prognoses of population persistence and evolutionary performance can be inferred only rarely from examination of vegetative characters alone.     

Comparative floral and vegetative differentiation between two European Aquilegia taxa along a narrow contact zone

As a first step in determining the identity and relative importance of the evolutionary forces promoting the speciation process in two closely related European taxa of Aquilegia, we investigated the levels of morphological variation in floral and vegetative characters over the narrow region where their ranges enter into contact, and evaluate the relative importance of both types of traits in their differentiation. A total of 12 floral and ten vegetative characters were measured on 375 plants belonging to seven A. vulgaris populations and six A. pyrenaica subsp. cazorlensis populations located in southeastern Spain. Floral and vegetative morphological differentiation occur between taxa and among populations within taxa, but only vegetative characters (particularly plant height and leaf petiolule length) contribute significantly to the discrimination between taxa. Differentiation among populations within taxa is mostly explained by variation in floral traits. Consequently, morphological divergence between the two taxa cannot be interpreted as an extension of among-population differences occurring within taxa. Multivariate vegetative, but not floral, similarity between populations could be predicted from geographical distance. Moreover, the key role of certain vegetative traits in the differentiation of A. vulgaris and A. p. cazorlensis could possibly be attributable to the contrasting habitat requirements and stress tolerance strategies of the two taxa. These preliminary findings seem to disagree with the currently established view of the radiation process in the genus Aquilegia in North America, where the differentiation of floral traits seems to have played a more important role.     

Cytogeography and meiotic chromosome configurations of six intraspecific aneuploids ofCarex conica Boott (Cyperaceae) in Japan

Chromosome numbers were determined for 340 plants ofCarex conica from 83 populations in Japan. Six aneuploids, 2n=32, 33, 34, 36, 37 and 38, were found. Plants with even diploid chromosome numbers 2n=32, 34, and 36 were the most common and had different geographical distributions. Individuals with 2n=32 were from islands in the Seto Inland Sea and nearby coastal areas of the Chugoku District of Honshu; those with 2n=34 were from the Kanto, Chubu and Kinki Districts of Honshu; those with 2n=36 were from the mountainous areas of Chugoku, Shikoku and Kyushu Districts. Canonical discriminant analysis of 17 morphological characters demonstrated that the plants with 2n=32 were clearly distinct from those with 2n=34 or 36. All four aneuploids with even chromosome numbers showed normal bivalent pairing at meiotic metaphase I and probably represent cytogenetically stable cytodemes. Plants with 2n=33 had one heteromorphic trivalent and 15 bivalents, indicating a structural mutation. At mitotic metaphase I, one chromosome was markedly larger than the others, suggesting that the 2n=33 plants arose from 2n=34 plants by fusion of two chromsomes. The plant with 2n=37 was intermediate in morphology betweenCarex conica (2n=36) andC. morrowii (2n=38) and probably originated as an interspecific hybrid between these species.     

Intraspecific evolution ofMicroseris pygmaea (Asteraceae,Lactuceae) analyzed by cosegregation of phenotypic characters (QTLs) and molecular markers (RAPDs)

The Chilean annual,Microseris pygmaea, has differentiated in distinct coastal and inland series of populations after long-distance dispersal from western North America. Two plants from the most diverse biotypes were crossed, a large F₂ was raised and analysed for segregation of 30 phenotypic characters. Segregation of molecular markers (47 RAPDs, 1 RFLP, 2 isozymes) was determined in a subpopulation of 45 plants which include all extremes for the phenotypic characters. 32 marker/character cosegregations were significant at the 1% level in t-tests between dominant and homozygous recessive marker genotypes. Considering linkage among markers and pleiotropy of certain marker loci, the number of independent quantitative trait loci (QTLs) is reduced to about 18. Interactions among 2 or 3 QTLs affecting one character have been characterized. The phenotypic differentiation ofM. pygmaea during its evolution from a single founder individual begins to be understood at the level of single-gene mutants.     

A phenetic study of Cassia sensu lato (Leguminosae-Caesalpinioideae: Cassieae: Cassiinae) in Thailand

Cassia L. sensu lato, a large heterogeneous genus of flowering plants, occurs naturally in the tropics around the world. Recent works based on floral morphology have supported a division of this genus into three genera, namely Cassia L. s. str., Chamaecrista Moench and Senna Mill. In order to investigate this new classification, 508 specimens of 18 taxa of the genus Cassia s.l. grown in Thailand were analyzed using cluster analysis and canonical discriminant analysis. The total 32 vegetative and reproductive morphological characters were employed in these analyses. In cluster analysis, Cassia s.l. can be separated into four groups, respectively viz. Chamaecrista, Senna alata, Senna and Cassia s /str. The four-cluster grouping is discussed. From a canonical discriminant analysis using the four-cluster grouping as a priori groups, it can be concluded that Cassia s. str., Senna, and Chamaecrista are indeed distinct taxa. The three most important characters that separate the three genera are filament length, fruit length, and ovary stalk length. These quantitative characters, together with some qualitative characters, were useful in constructing an identification key to these genera. Among the three genera, it was also found that Senna is rather a heterogeneous taxon. The difference between the studied species was discussed.     

Vegetative diversification and radiation in subtribeDendrobiinae (Orchidaceae): Evidence from chloroplast DNA phylogeny and anatomical characters

The data derived from a chloroplast DNA restriction site analysis of subtribeDendrobiinae (Orchidaceae) indicate that extreme vegetative diversification is concentrated in two limited parts of this group. Overlaying the vegetative character states onto the chloroplast DNA cladogram suggests that several xeromorphic, vegetative characters evolved in the lines leading to the above-mentioned clades. Several anatomical characters are also associated with xeromorphy. These vegetative and anatomical characters facilitated the establishment of this group in various dry habitats. On the other hand, the modifications of size and number of parenchymatous cells substantially contributed to the vegetative diversification. This fact implies that a simple structural adjustment can result in a major modification of growth habits in theDendrobiinae.     

Phenotypic plasticity inCrepis tectorum (Asteraceae)

Two experiments were carried out using two different approaches to compare populations ofCrepis tectorum (Asteraceae). One was based on a comparison of means of various vegetative and reproductive characters and another was based on a comparison of response patterns of the same characters in a series of environments. Population divergence within two earlier recognized form series, one from weed habitats and one from alvar habitats on Baltic islands, resulted in a partially overlapping pattern in cluster analyses based on character means. However, the pattern revealed by a comparison of the direction and amount of plastic response suggested that populations within the two form series had more similar response patterns than other combinations of populations. It was concluded that patterns of plasticity may provide useful additional information on the overall similarity among taxa. An hypothesis that plants in weed populations should exhibit a greater phenotypic response to the environments than plants in alvar populations was rejected.     

Somacional variation and eyespot toxin tolerance in sugarcane

The analysis of sugarcane plants regenerated from culture for their reaction to eyespot (Helminthosporium sacchari) toxin is described. A total of 480 culture-derived plants (somaclones) from cultivar Q101 were characterized. Some of these plants derived from cultures which had been subjected to selection with the eyespot toxin and others were derived without overt selection. Leaves were assayed for their toxin reaction. A very high frequency of toxin-tolerant variants was found. The distribution was even further biased toward resistance in those plants regenerated from cultures exposed to toxin selection.A total of 85 somaclones was analysed for the stability of their increased toxin tolerance to the primary somaclone; 22% were more tolerant; 38% were more susceptible. These results are discussed as they relate to the possibility of using consecutive vegetative segregation.Six tolerant variants were also passed through a second tissue culture cycle and 60 secondary somaclones were assayed. Twenty four (40%) of these plants had a similar tolerance to the primary somacione; 22% were more tolerant; 38% were more susceptible. These results are discussed as they relative to the possibility of using consecutive cycles of culture to stack improved characters into a sugarcane cultivar.     

The introduction and release of Chiasmia inconspicua and C. assimilis (Lepidoptera: Geometridae) for the biological control of Acacia nilotica in Australia

Two geometrid moths Chiasmia inconspicua and Chiasmia assimilis, identified as potential biological control agents for prickly acacia Acacia nilotica subsp. indica, were collected in Kenya and imported into quarantine facilities in Australia where laboratory cultures were established. Aspects of the biologies of both insects were studied and CLIMEX® models indicating the climatically favourable areas of Australia were developed. Host range tests were conducted using an approved test list of 74 plant species and no-choice tests of neonate larvae placed on both cut foliage and potted plants. C. inconspicua developed through to adult on prickly acacia and, in small numbers, Acacia pulchella. C. assimilis developed through to adult on prickly acacia and also in very small numbers on A. pulchella, A. deanei, A. decurrens, and A. mearnsii. In all experiments, the response on prickly acacia could be clearly differentiated from the responses on the non-target species. Both insects were approved for release in Australia. Over a three-year period releases were made at multiple sites in north Queensland, almost all in inland areas. There was no evidence of either insect’s establishment and both colonies were terminated. A new colony of C. assimilis was subsequently established from insects collected in South Africa and releases of C. assimilis from this new colony were made into coastal and inland infestations of prickly acacia. Establishment was rapid at one coastal site and the insect quickly spread to other infestations. Establishment at one inland area was also confirmed in early 2006. The establishment in coastal areas supported a CLIMEX model that indicated that the climate of coastal areas was more suitable than inland areas.     

Comparing the effects of the exotic cactus-feeding moth, Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae) and the native cactus-feeding moth, Melitara prodenialis (Walker) (Lepidoptera: Pyralidae) on two species of Florida Opuntia

This study examined the effects of the native cactus moth borer, Melitara prodenialis, and the invasive cactus moth borer, Cactoblastis cactorum, on two common cactus species, Opuntia stricta and O. humifusa at coastal and inland locations in central Florida. Opuntia stricta were present only at coastal sites and O. humifusa were present at coastal and inland sites. Throughout the duration of the study, coastal plants were subject to damage solely by C. cactorum and inland plants solely by M. prodenialis. Results showed marginally significantly higher numbers of eggsticks on O. stricta than O. humifusa and significantly higher numbers at coastal sites than at inland sites. There was also significantly higher moth damage on O. stricta than O. humifusa and at coastal sites than inland sites, but not significantly so. However, there was a higher level of plant mortality for O. humifusa than for O. stricta and a significantly higher level of cactus mortality at inland sites when compared to coastal sites. This increased mortality may be due to increased attack by true bugs, Chelinidea vittiger, and by Dactylopius sp., combined with attack by M. prodenialis. Inland plants also tended to be smaller than coastal plants and could be more susceptible to the combined effects of all insects. Further long-term research on coastal cactus survival when attacked and unattacked by Cactoblastis is necessary to fully determine the effects of this moth on Opuntia survival.     

Concordant clines and significant correlation between floral and pollen characters in Asarum heterotropoides var. heterotropoides (Aristolochiaceae)

To examine geographic differentiation in Asarum heterotropoides var. heterotropoides in Hokkaido Is. in the northern Japan, in which two putative cryptic species have been suspected to exist, extensive and detailed morphological research on 794 individuals from 44 populations throughout Hokkaido Is. was performed. Among the characters examined, the angle between and tip shape of the calyx lobes and the supratecta of the pollen grains were significantly correlated and were found in similar geographic clines. Among them, the pollen showed two discrete states in almost distinct distribution. Multidimensional scaling analysis showed that individuals within each of the two pollen types had different trends in flower characters. Consequently, we assumed a cline from south to north on Hokkaido Is. For the causes of the cline, the two hypotheses were proposed, primary geographic differentiation or extensive introgressive hybridization between two distinct geographical species existing in the past.     

夹竹桃科2种引种植物分泌结构的解剖学研究

为了解夹竹桃科(Apocynaceae)植物乳汁管的发生发育,对爱之蔓(Ceropegia woodii)和百万心(Dischidia ruscifolia)营养器官中的分泌结构进行了显微观察。结果表明,爱之蔓和百万心营养器官中均有无节分枝乳汁管的分布,茎皮层中的乳汁管大部分具有明显的分枝,叶中乳汁管具明显分枝,分布与走向多与叶脉维管组织平行。另外,爱之蔓营养器官中的分泌结构除乳汁管外,还有分泌腔。这为夹竹桃科植物的系统分类研究提供了解剖学依据。     

Nitrogen fertilizer and gender effects on the secondary metabolism of yaupon,a caffeine-containing North American holly

Females and males of sexually dimorphic species have distinct resource demands due to differential allocation to reproduction. Sexual allocation theory predicts that functional traits will diverge between sexes to support these demands. However, such dimorphism may be masked by the impact of current reproduction on source-sink interactions between vegetative and reproductive organs. We ask whether natural selection has led to genetic dimorphism in homologous physiological traits between sexes of the dioecious willow shrub, Salix glauca. In a common garden experiment we compared physiological responses to drought stress by male and female ramets in the absence of confounding demands from reproductive structures. Ramets experienced similar pre-dawn leaf water status (Ψ_l) as parental genets in flower within the natural population, indicating that experimental dry-down mirrored environmental conditions in nature. Male and female ramets achieved similar instantaneous water use efficiency, based on the ratio of carbon gain to water loss, under wet and dry conditions. However, female ramets experienced greater water stress (i.e., more negative Ψ_l) than males under dry conditions. Lower Ψ_l for female ramets may partly reflect the maintenance of conductance under drought; males, in contrast, maintain Ψ_l under drought by reducing conductance. Differences between sexes in terms of conductance and leaf water status of the vegetative ramets were absent in a concomitant comparison of parental flowering plants. Our results show (1) genetic divergence in physiology between sexes of S. glauca occurs in the absence of gender-specific reproductive sinks, (2) males are the more physiologically plastic sex with respect to water use, and (3) paradoxically, divergence in water relations between sexes is not detectable at sexual maturity under natural conditions.     

The genusAntennaria (Asteraceae: Inuleae) in western North America: Morphometric analysis ofAntennaria alborosea,A. corymbosa,A. marginata,A. microphylla,A. parvifolia,A. rosea,andA. umbrinella

Multivariate analysis of vegetative and reproductive characters was used to examine morphological relatedness amongAntennaria alborosea A. E. Porsild,A. corymbosa E. Nels.,A. marginata Greene,A. microphylla Rydb.,A. parvifolia Nutt.,A. rosea Greene, andA. umbrinella Rydb. Both pistillate and staminate plants were examined. Some of the characters examined were variable in one species, but stable in another (i.e., presence or absence of papillae on the achenes). Our analyses indicate that the seven species are morphologically distinct. It is hypothesized that theA. rosea agamic complex arose through hybridization amongA. corymbosa, A. microphylla, A. umbrinella, and possiblyA. dioica (L.)Gaertn. However, hybridization between the three former species and others, as well as their subsequent morphological responses to different environmental conditions causes confusion in recognizing the taxa.Antennaria angustifolia Rydb.,A. arida E. Nels.,A. confinis Greene,A. scariosa E. Nels.,A. foliacea humilis Rydb.,A. concinna E. Nels., andA. viscidula E. Nels. are considered to represent F 1 hybrids.     

Tissue culture-induced DNA methylation polymorphisms in repetitive DNA of tomato calli and regenerated plants

The propagation of plants through tissue culture can induce a variety of genetic and epigenetic changes. Variation in DNA methylation has been proposed as a mechanism that may explain at least a part of these changes. In the present study, the methylation of tomato callus DNA was compared with that of leaf DNA, from control or regenerated plants, at MspI/HpaII sites around five middle-repetitive sequences. Although the methylation of the internal cytosine in the recognition sequence CCGG varied from zero to nearly full methylation, depending on the probe used, no differences were found between callus and leaf DNA. For the external cytosine, small differences were revealed between leaf and callus DNA with two probes, but no polymorphisms were detected among DNA samples of calli or DNA samples of leaves of regenerated plants. When callus DNA cut with HindIII was studied with one of the probes, H9D9, most of the signal was found in high-molecular-weight DNA, as opposed to control leaf DNA where almost all the signal was in a fragment of 530 bp. Also, an extra fragment of 630 bp was found in the callus DNA that was not present in control leaf DNA. Among leaves of plants regenerated from tissue culture, the 630-bp fragment was found in 10 of 68 regenerated plants. This 630-bp fragment was present among progeny of only 4 of these 10 plants after selfing, i.e. it was partly inherited. In these cases, the fragment was not found in all progeny plants, indicating heterozygosity of the regenerated plants. The data are interpreted as indicating that a HindIII site becomes methylated in callus tissue, and that some of this methylation persists in regenerated plants and is partly transmitted to their progeny.     

The pleiotropic effects induced by therolC gene in transgenic plants are caused by expression restricted to protophloem and companion cells

Expression of therolC gene fromAgrobacterium rhizogenes causes morphological and developmental alterations in transgenic plants. The histological alterations underlying the macroscopic changes and the cellular localization of the site of expression of therolC gene have shown that: (i) the expression of therolC gene is developmentally regulated, (ii) in vegetative transgenic plants, the expression of therolC gene under the control of its own promoter is restricted to companion and protophloem cells, (iii) the site of action of the product(s) of the activity of the rolC enzyme is distinct from its site of expression, (iv) precise localization of the rolC peptide has been achieved by immunocytochemistry but not by the histochemical GUS assay. These results imply that the sites of action and expression of therolC gene in trangenic plants are physically separated. Thus the product(s) of the activity of the rolC enzyme must be a factor capable of being transported. Current models forrolC gene action are discussed taking into account the reported results.     

Population dynamics of an ice-associated diatom, Thalassiosira australis Peragallo, under fast ice near Syowa Station, East Antarctica, during austral summer

A clear shift from vegetative cells to auxospores and resting spores in Thalassiosira australis was observed in the water column and sinking fluxes under the fast ice near Syowa Station in the austral summer of 2005/2006. This is the first report of the auxosporulation by T. australis in situ. Resting spores were also observed in the sediment even before new spore formation, suggesting that T. australis can overwinter in the sediment. Heterotrophic dinoflagellates ingested and digested vegetative cells and auxospores but did not digest resting spores, suggesting a high tolerance of resting spores to grazing by heterotrophic dinoflagellates. We discuss the possible life history and overwintering strategy that T. australis uses in an Antarctic coastal area to cope with the unpredictable timing of sea ice growth and decay.     

Effects of simulated herbivory in three old field Compositae with different inflorescence architectures

The effects of simulated herbivory (early or late defoliation and cutting of the flowering shoot) on the growth and reproduction of three species of monocarpic composite forbs (Crepis pulchra, Picris hieracioides and C. foetida) with different inflorescence architectures were studied in experimental plots. For the three species studied, early defoliation had no significant effect on subsequent growth. In contrast, late defoliation, occurring at the start of the season of drought, had a negative effect on growth and reproduction in the two Crepis species, particularly C. foetida, but had less effect on P. hieracioides. Sexual biomass was more clearly affected by late defoliation than was vegetative biomass, although the effects differed markedly among species possibly as a result of differences in phenology. Clipping the flowering shoot removed about 3 times less biomass than late defoliation and had little effect on vegetative biomass. It had much greater effects on the sexual biomass in P. hieracioides and C. pulchra, and resulted in the production of many shoots sprouting from the rosette, allowing the treated plants to regain a vegetative biomass close to that of control plants. Clipping did however lead to the production of shorter shoots and a reduction in the number of capitula formed. In C. foetida, much branching occurred even when the main shoot was not cut; the architecture of individual plants was therefore only slightly changed by clipping the apical bud and the sexual biomass of this species was not affected by ablation of the flowering shoot. Overcompensation was found in only two families of C. pulchra for vegetative biomass. No over-compensation was found for sexual biomass, despite an increase in the number of flowering shoots in C. pulchra and P. hieracioides following clipping. However situations close to compensation for the vegetative biomass in the three species and in P. hieracioides for the sexual biomass were recorded. The response of the three study species to simulated herbivory were related to their architecture and to the time of defoliation.     

Use of monocharacteristic growth forms and phenological phases to describe and differentiate plant communities in Mediterranean-type ecosystems

The ecomorphological and phenological study was carried out within aMediterranean vegetation context, in Quercus suberforests,which have been substituted by shrublands of Cistus sppwithin two Natural Parks in the south of the Iberian Peninsula. Theecomorphological characters that show meaningful differences between both typesof vegetation are: location of renewal buds, spinescence, stratification,maximum height of the vegetation, organs periodically shed, leaf consistency,leaf tomentosity, leaf size, and life duration of leaves, plant duration,vegetative regeneration after fire, main vegetative growth season, mainflowering season and fruit type. The phenological phases also help to discernbetween forest and shrubland, specially flower bud formation, fruiting, seeddispersal, and the existence of brachyblast vegetative growth and brachyblastleaf shedding. We propose three new indexes based on phenological phases:active period of the species (APS), active period of thecommunity (APC) and reproductive/vegetative activity of thespecies (RVA). The results of their application, in combination with theecomorphological characters, have proved promising in describing vegetation andin clearly differentiating communities. The results also show the existence ofdifferent ecomorphological groups of plants at community level, with consequentecological, historical, phytocoenological and adaptive implications