Sexual differences in biomass and nutrient allocation of first-year Silene dioica plants

Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences. Received: 07 May 1998 / Accepted: 30 October 1998     

In vitro organogenesis in two dioecious species,Rumex acetosella L. and R. acetosa L. (Polygonaceae)

Callus cultures were established from dioecious plant species Rumex acetosella and R. acetosa, using cotyledons, hypocotyls and stem tips of aseptically germinated seedlings as primary explants. Cultures were also established from male and female R. acetosella adult plants, starting from vegetative lateral buds. Cell division was induced using a high 2,4-D concentration, while bud induction and multiplication were stimulated on a medium with high BAP/IAA ratio. Cotyledon fragments of both species produced only rhizogenic calli. Hypocotyl-derived calli of R. acetosella produced buds, while those of R. acetosa showed no bud forming response under these conditions. Bud multiplication occurred in stem tip cultures of both species and in lateral bud cultures of R. acetosella. Calli derived from male plants produced more buds than those from female. Shoots were easily rooted using IBA, and plantlets were effectively transferred to soil. Flowering was not induced in culture. The sex of regenerated male and female plants was not altered by the culture conditions.     

Rumex subgen.Rumex sect.Axillares (Polygonaceae) in South America

Rumex sect.Axillares first described by the author in 1937 is known to be represented in N. America by 20 endemic species. From S. America only five species, all endemic, were previously known; this number is raised in the present revision to 15. Outside America there exist small groups of endemic species in S. Africa and in the islands of Hawaii as well as in Australia and scattered single species in C. Africa, on the remote Atlantic islands of Tristan de Cunha and Gough Island, and in eastern Asia.—In Europe the section is completely absent, except for three American species which occur as rare aliens. TheAxillares are not only morphologically well-characterized by their growth—no leaf rosettes, flowering period practically unlimited by formation of later flowering axillary shoots, flowers often monoecious—but also by their genetic structure expressed by a different behaviour in hybridization. Hybrids are frequent between species of sect.Rumex (syn.Simplices) and are easy to recognize by their sterility which results in a certain peculiarity in habit. In contrast, no spontaneous hybrids are known with certainty among species of sect.Axillares. Most artificial hybrids are fairly fertile. So far only three hybrids between species of the two sections have been known. Three more have been found by the author in Chile, all being highly sterile and showing the growth characteristics resulting from sterility as occur in hybrids between species of sect.Rumex.

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Resumen Rumex secciónAxillares fue descrito por primera vez por el autor en 1937. Se conoce en América del Norte con 20 especies endémicas. De América del Sur se conocían sólo 5 especies, todas ellas endemicas; este número asciende a 15 en la presente revision. Fuera de las Américas se encuentran pequeños grupos de especies endémicas en Sudáfrica, en las Islas de Hawaii y en Australia; especies aisladas se encuentran en Africa Central, en las lejanas Islas Atlánticas de Tristan de Cunha y Gough Island y en Asia Oriental. La sección no tiene representantes en Europa, con excepción de tres especies americanas introducidas, poco frecuentes. La secciónAxillares se caracteriza bien, tanto morfológicamente por su forma de creciemiento—sin rosetas foliares, período floral prácticamente ilimitado por la formación de inflorescencias axilares parciales, flores a menudo monoicas—como por su estructura genética que se manifiesta en un comportamiento diferente de hibridación. Híbridos son frecuentes en las especies de la secciónRumex (syn.Simplices) y reconocibles por su esterilidad que se expresa en cierta particularidad del habito. Al contrario, no hay seguridad de híbridos espontáneos en la secciónAxillares. La mayor parte de los híbridos artificiales son totalmente fértiles. Hasta ahora se conocían solo tres híbridos entre las especies de ambas secciones. Otros tres han sido encontrados por el autor en Chile. Todos completamente estériles y con las características de hábito típicas resultantes la esterilidad, como ocurre en los híbridos de la secciónRumex.

     

Induction of somatic embryogenesis and embryo development in Rumex acetosella L.

Axillary buds of the dioecious plant Rumex acetosella L. were isolated and cultured in vitro. The callus tissue which developed at the basal parts of the explants displayed a high capacity for shoot formation. This morphogenetic pattern was predominant on Murashige and Skoog (MS) medium supplemented with 2% sucrose, 2.2 mgl^-1 benzylaminopurine and 0.17 mgl^-1 indole-3-acetic acid. Somatic embryogenesis was induced when the osmolality of the medium was increased by adding 6% sucrose instead of 2%, or hexitols in addition to 2% sucrose. Most of the embryogenic calli were formed on the basal parts of leaf laminae and bracts. Development and maturation was strongly promoted by transferring the tissue to a solid or liquid medium lacking benzylaminopurine and indole-3-acetic acid and supplemented with 10 mgl^-1 gibberellic acid. The embryos germinated and developed into normal rosette plants when transferred to vermiculite moistened with hormone-free, half-strength MS salt solution. The histology of successive embryogenic stages is presented.     

Responses to moisture stress in male and female plants of Rumex acetosella L. (Polygonaceae)

Summary Male and female plants of Rumex acetosella were grown on a moisture gradient to measure possible differences in the drought tolerance of the sexes. The growth of both sexes declined under water stress but males were significantly more drought tolerant. This could not be explained by greater water use efficiency in the male plants; measured rates of both photosynthesis and leaf conductance did not differ significantly between the sexes. Multiple discriminant analysis showed that the sexes differed at all moisture regimes in their overall patterns of biomass allocation. Males had proportionately greater investment in root and leaf tissue which could explain their growth advantage over females under water stress. Despite essentially equal water use efficiencies, on a per plant basis males, with more leaf and root biomass, could fix more carbon and more rapidly exploit the local water resource than females. Thus the pattern of biomass allocation rather than intrinsic physiological differences appears to explain the greater drought tolerance of male plants of Rumex acetosella.     

Effects of cadmium on integration and resource allocation in the clonal fern Salvinia molesta

Summary In clonal plants exposed to pollution, ramets which are physiologically integrated may be less fit than ramets which are independent, if (a) translocation of toxins from contaminated ramets produced toxicity, or (b) toxicity in parent ramets reduced the degree of nutritional support to otherwise non-exposed daughters. These hypotheses were tested in the aquatic fern Salvinia molesta exposed to cadmium. Pre-treatment of parent ramets with cadmium decreased the number and biomass of daughters subsequently produced in a cadmiumfree medium, because of reduced parental support of the first daughter generation. Second generation and later daughters were unaffected. Pre-treatment did not affect the pattern of integration (which, in terms of apical daughters' biomass, was bimodal with increasing colony size), or concentrations of essential elements in new growth. However, a diversion of resources from lateral to apical daughters occurred as a result of pre-treatment, especially in colonies with 3 attached parents. Loss/gain analysis showed that the diversion was almost reciprocal in terms of biomass, ramet numbers and phosphorus content. Integration between contaminated and uncontaminated ramets was not disadvantageous to the clone as a whole. However, integration was disadvantageous for Ca, Mg and Zn concentrations in daughters, which declined 15–22%. Because of enhanced apical growth, an indirect benefit of integration may be a more rapid fragmentation and dispersal of daughters from the site of contamination than if the parents were independent.     

Sex ratio variation and spatial segregation of the sexes in populations ofRumex acetosa andR. acetosella (Polygonaceae)

Populations ofRumex acetosa andR. acetosella were studied during two growth seasons. The ramet sex ratios ofR. acetosa were always female-biased. InR. acetosella the sex ratios expressed more variation but were mostly female-biased. In both species the sex ratios commonly varied between subpopulations reflecting a partial spatial segregation of the sexes. No marked differences between sexes in vegetative vigour were detected in either species. Interactions between sex ratios, various soil characteristics and population densities were determined. Possible mechanisms for causing biased sex ratios and partial spatial segregation of the sexes are discussed.     

The comparative germination ecology of nine Rumex species

The germination requirements, dormancy cycle and longevity of nine Rumexspecies were studied in field conditions and laboratory experiments to show theadaptations of the related species to their specific habitat. Within one genus,rather striking differences were observed in germination ecology. However, theclosely related species, R. acetosa and R.scutatus, are very similar: they fruit in early summer; theirseeds can germinate immediately after dispersal, and they are nondormant andshort-lived. R. acetosella also has fruits insummer, but the seeds do not germinate the first season after dispersal. Theyare long-lived, but buried seeds do not show a dormancy cycle; they mightgerminate in different seasons after exposure to light. Seeds of four species (R. conglomeratus,R. maritimus, R. sanguineus andR. crispus) are long-lived and undergo aseasonal dormancy cycle, with a low level of dormancy in winter and early springand a deep dormancy in summer as was already known for R.obtusifolius. These seeds are shed in the autumn, and they germinatemainly in the spring in consecutive years. R. maritimusalso germinates in summer and autumn on drying muddy soils. The seeds of R. hydrolapathum only germinate onwaterlogged soils, which explains its growth at the edge of streams and ponds.Its seeds are rather short-lived. The seeds of the species on very wetplaces require a higher temperature for germination.     

Size and sex allocation in monoecious woody plants

The female size advantage hypothesis predicts that the allocation ratio of female: male reproductive effort should increase with plant size (total reproductive effort). A male height advantage hypothesis has also been proposed, based on the supposed greater advantage of height to male reproductive success in wind-pollinated plants. These ideas were tested with data for wind-pollinated, monoecious trees and shrubs which exhibit a suitably large range of sizes. Number of male inflorescences increased faster with size than did number of female inflorescences in 2 of 9 species; in the remaining 7 species there was no significant difference. The male:female ratio of inflorescence numbers increased with height in 4 of 7 species and did not change significantly in the remaining 3 species, as shown by regression. Height and size are highly correlated and so their effects could not be distinguished. The fact that many conifers place the female cones uppermost in the crown suggests that size and not height favors increased allocation to male function, as does well-established theory connecting the existence of male versus female size advantage to pollen and seed dispersal chacteristics. Regression analysis of the relation between male and female reproductive effort should be done by reduced major axis regression; ordinary least squares regression underestimates slopes; in this study opposite conclusions could be drawn from ordinary least squares and reduced major axis regressions.     

Theory of resource allocation strategy in territorial male tropical reef fishes

A mathematical model is proposed to explain energy resource allocation between sperm production and territoriality in male reef fishes (Labridae species) from the point of view of optimization. Labridae species are typically characterized by both TP (terminal phase) and IP (initial phase) males. The former are considered to release a lower amount of sperm but show aggressive territoriality. In the model, TP male reproductive success is considered as depending upon both fertilization probability (depending on sperm density) and the individual's own territorial activities. Between these factors, a trade-off exists by which the fertilization probability can be enhanced only by reducing territoriality. Therefore, the male has to decide how much of the total available energy resource should be allocated to each. The model showed that under high fertilization efficiency the male can achieve high success by spending less of the resource on sperm production and correspondingly more for territoriality. The TP male reproductive success increases with decreasing male density in the habitat. Nevertheless, when intruding males cannot be excluded completely by territorial behavior of the TP male, females prefer high male density. If females can control the number of intruding males to some degree, conflict may arise between the sexes. Received: December 8, 1999 / Accepted: May 24, 2000     

Characterization of PSII photochemistry and thermostability in salt-treated Rumex leaves

A study was conducted, using chlorophyll fluorescence, rapid fluorescence induction kinetics, and polyphasic fluorescence transients, to determine the effect of salt treatment and heat stress on PSII photochemistry in Rumex leaves. Salt treatment was accomplished by adding NaCl solutions of different concentrations ranging from 50 to 200 mmol/L. Heat stress was induced by exposing the plant leaves to temperatures ranging from 29 to 47 degrees C. The control plants were grown without NaCl treatment. The data acquired in this study showed that NaCl treatment alone had no effect on the maximal photochemistry of PSH or the polyphasic rise of chlorophyll fluorescence. However, the NaCl treatment modified heat stress on PSII photochemistry in Rumex leaves, which was manifested by a lesser heat-induced decrease in photochemical quenching (qP), efficiency of excitation energy capture by open PSII reaction centers (Fv'/Fm'), and quantum yield of PSII electron transport (phiPSII). The data also showed that NaCl treatment compromised the impact of heat stress on the capacity of transferring electrons from Q(A)- to Q(B). Furthermore, the NaCl treatment promoted heat resistance of O2-evolving complex (OEC). In summary, NaCl treatment enhanced the thermostability of PSII.     

Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae)

Abstract Patterns of reproductive and vegetative biomass allocation were compared in male and female plants of the alpine herb Aciphylla simplicifolia. Male and female plants had similar vegetative biomass but differed in the pattern of resource allocation. Inflorescences of males and females were similar in weight at the time of flowering, but differed in biomass allocation to some structures within the inflorescences, particularly those associated with ovule production and pollinator attraction (number and size of flowers). At the time of fruit production, female inflorescences were 2.6 times heavier than at flowering with developing fruit six times heavier than flowers. In addition to the increase in biomass allocated to structures associated with the provisioning and dissemination of seed, support structures (main and side stalks) were also heavier. As a result of this additional investment of resources at the time of fruit production, the reproductive effort (RE) of female plants was much higher than that of males: 37% of above ground biomass compared with 21% for males. Differences in RE did not change with plant size; however, allocation to reproduction appeared to be a constant proportion of biomass over nearly all plant sizes sampled. These results show that sex‐specific resource allocation can be a complex of temporal and morphological patterns.     

Endogenous cytokinins in the vegetative and reproductive phases of development in the dioecious plant Rumex acetosella L.

The content of endogenous cytokinins has been analysed in leaf and inflorescence extracts of male and female R. acetosella plants, using gas chromatography. Plant parts were extracted at four stages of development: leaves of juvenile plants, leaves of adult plants at the time of flower initiation and in full bloom, and upper internodes of the inflorescence stalks. Cytokinins with characteristics similar to isopentenyl adenine and adenosin, zeatin, zeatin riboside, and a bound form of zeatin, were all found in the extracts. The total amount of cytokinins was higher in female than in male plants during all these stages.Reprint requests     

Effects of herbivory by chrysomelid beetles on the growth and survival ofRumex plants

The effects of herbivory by the guild of chrysomelid beetles on the growth and survival ofRumex plants were examined in relation to the distribution and size of the plants. Gastrophysa atrocyanea never appeared on solitary plants whereasMantula clavareaui showed even utilization of solitary and clumped plants.Galerucella vittaticollis utilized large solitary plants most frequently. In patches of the host plants, the frequency of withering of the aerial parts was higher on small plants. The regrowth rate was higher when the aerial parts withered in spring than in summer. In patches of the host plants, the regrowth rate was higher on small plants. On the other hand, it was lower on small solitary plants than on large solitary plants and small plants in patches. In large plants, no difference in survival rate was recognized between solitary and clumped plants, because both the frequency of withering of the aerial parts and the regrowth rate were lower in clumped than solitary plants. The high vigor against intense herbivory by the chrysomelid beetles brought on the delay in the phenology ofRumex plants. This shift permits in parts the existence ofG. vittaticollis in mid summer. The temporal and spatial interactions between herbivore guild and the host plants were discussed.     

The effects of chrysomelid beetle grazing and plant competition on the growth of Rumex obtusifolius

Summary The interaction between grazing by Gastrophysa viridula and interspecific plant competition was investigated for Rumex obtusifolius growing in the field. During an eight-month growing season non-competing R. obtusifolius grew larger than competing plants although herbivore loads (beetle productivity per unit leaf area) were similar for competing and non-competing plants. However, grazing significantly reduced the growth of R. obtusifolius only in competing plants thus demonstrating a synergism between plant competition and invertebrate herbivory. The relevance of these findings for the distribution of plant species is discussed.     

Elimination of oxalate contamination in RNA isolation from<Emphasis Type="Italic">Rumex obtusifolius</Emphasis>

Many plant RNA isolation techniques aim to prevent contamination by means of secondary phenolics, carbohydrates, RNase, and other chemicals. However, when applied in our laboratory to the isolation of RNA fromRumex obtusifolius, these protocols failed to produce good quality RNA. A major problem was contamination of the RNA samples with the secondary metabolite oxalate. The relative quantities of guanidine isothiocyanate extraction buffer to plant tissue used in the protocol had significant effects on oxalate contamination. An increase in extraction buffer, from 1.5 mL in the original method to 15 mL per 200–300 mg of tissue in our protocol, removed the oxalate from the RNA. This RNA was of a good quality and was suitable for molecular biology applications.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Interaction of light and temperature on seed germination of Rumex obtusifolius L.

Germination of Rumex obtusifolius L. seeds (nutlets) is low in darkness at 25° C. Germination is stimulated by exposure to 10 min red light (R) and also by a 10-min elevation of temperature to 35° C. A 10-min exposure to far-red light (FR) can reverse the effect of both R (indicating phytochrome control) and 35° C treatment. Fluence-response curves for this reversal of the effect of R and 35° C treatments are quantitatively identical. Treatment for 10 min with light of wavelenght 680, 700, 710 and 730 nm, after R and 35° C treatment, demonstrates that germination induced by 35° C treatment results from increased sensitivity to a pre-existing, active, far-red-absorbing form of phytochrome (Pfr) in the seeds.Abbreviations FR far-red light - P phytochrome - Pr red-absorbing form of P - Pfr far-red-absorbing form of P - R red light     

Ecophysiological differences between male and female plants of Pistacia lentiscus L.

Previous studies in spatial distribution of male and female shrubs of Pistacia lentiscus have demonstrated that less perturbed areas, older communities with a well developed cover, have male-biased sex ratios, whereas in abandoned old agricultural areas there are no significant differences between the number of male and female plants. In this study, we analyse both sexes in terms of their photosynthetic features that could provide a physiological basis for habitat partitioning between sexes. Rates of light-saturated assimilation and stomatal conductance were studied in male and female plants during summer. Assimilation rates were higher in the morning than in the afternoon and mean daily maximum assimilation rates reached 10.9 and 6.6 mol m^–2 s^–1, for male and female plants, respectively. In the absence of drought stress (laboratory conditions), the measured photosynthetic characteristics of leaves of male and female plants, provided by fluorescence studies and light and CO₂ response curves, were similar. Under natural stress conditions however, lower CO₂ assimilation rates and stomatal conductances were recorded in female plants. The differences in the light response curve of effective quantum yield (_II) recorded under stress conditions showed also higher quantum yield for male plants under low irradiances. From this study we suggest that the differences observed between male and females are largely due to different degrees of stomatal control rather than to differences in photosynthetic activity, leading to higher water use efficiency (WUE) in female plants. However, despite the higher leaf control of water loss by females, they reduce the water potential to the same values as male plants, probably due to specific characteristics of the root system or of the conducting xylem. These results suggest that the ecological advantage of male plants in older communities is due to a higher competition for water uptake, while in the youngest open areas is the higher WUE in female plants that confer an ecological advantage.     

The influence of topography on male and female fitness components of Atriplex canescens

The influence of environmental heterogeneity on components of male and female fitness is examined using Atriplex canescens growing on steep slopes and alluvium at the slope base as a model system. Female fitness is estimated as the grams of fruit produced per plant and the grams of fruit per gram leaf tissue. Male fitness is estimated as the grams of stamens produced, the number of pollen grains dispersed to a given distance, and the potential number of grams of fruit sired taking into consideration the number and distribution of mates and competing pollen donors. The influence of increased plant size on male and female fitness components, the cost of reproduction (as measured by biomass, joules and nitrogen) on a gross level and a per offspring basis are also examined. The results indicate that the female function is more limited on the slope than the male function. The efficiency of pollen dispersal (the number of pollen grains per unit donor plant volume dispersed to a given distance) is enhanced by growing on slopes. Males become less efficient at dispersing pollen as they increase in size, while the efficiency of female reproduction (grams of fruit per volume or gram leaf tissue) is unaffected by increasing plant size. The cost to a male of siring a gram of fruit is about the same as the cost to a female of producing the gram of fruit. Implications for the evolution of sexual lability and dioecy are discussed.