Gender-specific costs of reproduction on vegetative growth and physiological performance in the dioecious shrub Corema album

Background and Aims

Reproductive costs imply trade-offs in resource distribution at the physiological level, expressed as changes in future growth and/or reproduction. In dioecious species, females generally endure higher reproductive effort, although this is not necessarily expressed through higher somatic costs, as compensatory mechanisms may foster resource uptake during reproduction.

Methods

To assess effects of reproductive allocation on vegetative growth and physiological response in terms of costs and compensation mechanisms, a manipulative experiment of inflorescence bud removal was carried out in the sexually dimorphic species Corema album. Over two consecutive growing seasons, vegetative growth patterns, water status and photochemical efficiency were measured to evaluate gender-related differences.

Key Results

Suppression of reproductive allocation resulted in a direct reduction in somatic costs of reproduction, expressed through changes in growth variables and plant physiological status. Inflorescence bud removal was related to an increase in shoot elongation and water potential in male and female plants. The response to inflorescence bud removal showed gender-related differences that were related to the moment of maximum reproductive effort in each sexual form: flowering in males and fruiting in females. Delayed costs of reproduction were found in both water status and growth variables, showing gender-related differences in resource storage and use.

Conclusions

Results are consistent with the existence of a trade-off between reproductive and vegetative biomass, indicating that reproduction and growth depend on the same resource pool. Gender-related morphological and physiological differences arise as a response to different reproductive resource requirements. Delayed somatic costs provide evidence of gender-related differences in resource allocation and storage. Adaptive differences between genders in C. album may arise through the development of mechanisms which compensate for the cost of reproduction.     

Sexual polymorphism,growth, and reproductive effort in Egyptian Thymelaea hirsuta (Thymelaeaceae)

Gender phenotypes of Thymelaea hirsuta (L.) Endl. were surveyed in six major habitats in the western Mediterranean region of Egypt. Five gender phenotypes were observed at all sites: subandroecious (male), subgynoecious (female), protogynous, protandrous, and gender-labile individuals. Patterns of vegetative growth, reproductive effort, and sex-size relationships were also determined. Females and males were comparable in overall abundance (35.5% and 30.2%, respectively). However, the frequency of sex forms varied significantly among habitats. Plant size (canopy volume) was used as a measure of environmental quality for the species: the coastal dunes were by far the most favorable habitat, followed by the nonsaline depressions, inland plateau, and inland siliceous deposits. The smallest plants were associated with the inland ridges and saline depressions. Protandrous individuals, and female shrubs at the more favorable habitats (e.g., the coastal dunes and nonsaline depressions), had greater canopy volume than males. In contrast, males were larger than females at the less favorable habitats (e.g., the inland plateau and saline depressions). Advantageous growth conditions at the coastal dunes are demonstrated by the greater rate of increase in crown diameter and crown volume noted there, for each size class and sex form. The greatest rates of increase were present in smaller-sized individuals. Size differences between the sex forms were considered in terms of trade-offs between individual growth rate (GR) and reproductive effort (RE); evidence of a trade-off was mixed. Indeed, in a comparison of GR and RE at two extremes of habitat (coastal dune vs. inland plateau), females had greater GR and greater RE at the relatively benign coastal dunes, while at the more harsh, inland plateau site males showed greater GR and RE than females. Peak reproductive effort varied significantly with gender phenotype and habitat type.     

Varietal influence of intercropped cowpea on the growth,yield and water relations of maize

The effects of three cowpea varieties namely, NewEra (a spreading type), Adzuki (an erect type) and Ife Brown (semi-erect) intercropped with maize on growth, yield and water relations of maize were investigated in the greenhouse and in the field. Cowpea varietal effects were significant on soil water extraction but insignificant on the performance of maize.     

Drought susceptibility of two tomato (Lycopersicum esculentum) varieties

The effects of three levels of moisture under greenhouse conditions and also identical moisture levels under field conditions, on the growth yield and water relations of two tomato cultivars, Fireball and Ife I, were investigated. The objective was to ascertain the drought susceptibility of these two tomato cultivars. The cultivar, Ife I, was more drought susceptible than Fireball and the drought susceptibility increased with the level of soil water stress. The drought tolerance, of Fireball is attributed to a lower leaf area, better root system development in terms of average root length and rooting depth, and a higher leaf water potential. The higher leaf stomatal resistances of Fireball variety suggest an inbuilt mechanism to regulate water vapour flow in times of stress.     

Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae)

Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998     

Effects of nitrogen nutrition and root medium water potential on growth, nitrogen uptake and osmotic adjustment of rice

The effects of nitrogen (N) nutrition on growth, N uptake and leaf osmotic potential of rice plants (Oryza sativa L. ev. IR 36) during simulated water stress were determined. Twenty-one-day-old seedlings in high (28.6 × 10 ^?4M) and low (7.14 × 10 ⁴M) N levels were exposed to decreased nutrient solution water potentials by addition of polyethylene glycol 6000. The roots were separated from the solution by a semi-permeable membrane. Nutrient solution water potential was ?0.6 × 10⁵ Pa and was lowered stepwise to ?1 × 10⁵, ?2 × 10⁵, ?4 × 10⁵ and ?6 × 10⁵ Pa at 2-day intervals. Plant height, leaf area and shoot dry weight of high and low nitrogen plants were reduced by lower osmotic potentials of the root medium. Osmotic stress caused greater shoot growth reduction in high N than in low N plants. Stressed and unstressed plants in 7.14 × 10⁴M N had more root dry matter than the corresponding plants in 28.6 × 10⁴M N. Dawn leaf water potential of stressed plants was 1 × 10⁵ to 5.5 × 10⁵ Pa lower than nutrient solution water potential. Nitrogen-deficient water-stressed plants, however, maintained higher dawn leaf water potential than high nitrogen water-stressed plants. It is suggested that this was due to higher root-to-shoot ratios of N deficient plants. The osmotic potentials of leaves at full turgor for control plants were about 1.3 × 10⁵ Pa higher in 7.14 × 10^?4M than in 28.6 × 10^?4M N and osmotic adjustment of 2.6 × 10⁵ and 4.3 × 10⁵ Pa was obtained in low and high N plants, respectively. The nitrogen status of plants, therefore, affected the ability of the rice plant to adjust osmotically during water stress. Plant water stress decreased transpiration and total N content in shoots of both N treatments. Reduced shoot growth as a result of water stress caused the decrease in amount of water transpired. Transpiration and N uptake were significantly correlated. Our results show that nitrogen content is reduced in water-stressed plants by the integrated effects of plant water stress per se on accumulation of dry matter and transpiring leaf area as well as the often cited changes in soil physical properties of a drying root medium.     

Causes of secondary sexual differences in plants — Evidence from extreme leaf dimorphism in Leucadendron (Proteaceae)

In extreme cases leaves in male plants of the dioecious genus Leucadendron (Proteaceae) are up to an order of magnitude smaller than female leaves. This secondary sexual dimorphism (SSD) in leaf size has previously been suggested to be due to intra-male sexual selection, leading to an increase in male allocation to reproduction in dimorphic species. After critically evaluating previous data provided to support this hypothesis, I suggest on both theoretical grounds and on re-analysis that this argument is unlikely and unsupported. Leaf size dimorphism could theoretically evolve directly due to disruptive ecological selection between genders, leading to niche dimorphism either within or between habitats. I test this ecological causation hypothesis by providing data on specific leaf area (sla) and water use efficiency (δ ¹³C) of leaves from males and females of several Leucadendron species. Results confirm the expectation of minimal gender differences. I argue that leaf dimorphism is a consequence of selection on flower size and architecture.     

Salt-induced modulation in growth,photosynthesis and antioxidant system in two varieties of Brassica juncea

The present study was carried out to examine salt-induced modulation in growth, photosynthetic characteristics and antioxidant system in two cultivars of Brassica juncea Czern and Coss varieties (Varuna and RH-30). The surface sterilized seeds of these varieties were sown in the soil amended with different levels (2.8, 4.2 or 5.6 dsm⁻¹) of sodium chloride under a simple randomized block design. The salt treatment significantly decreased growth, net photosynthetic rate and its related attributes, chlorophyll fluorescence, SPAD value of chlorophyll, leaf carbonic anhydrase activity and leaf water potential, whereas electrolyte leakage, proline content, and activity of catalase, peroxidase and superoxide dismutase enzymes increased in both the varieties at 30 d stage of growth. The variety Varuna was found more resistant than RH-30 to the salt stress and possessed higher values for growth, photosynthetic attributes and antioxidant enzymes. Out of the graded concentrations (2.8, 4.2 or 5.6 dsm⁻¹) of sodium chloride, 2.8 sm⁻¹ was least toxic and 5.6 dsm⁻¹ was most harmful. The variation in the responses of these two varieties to salt stress is attributed to their differential photosynthetic traits, SPAD chlorophyll value and antioxidant capacity, which can be used as potential markers for screening mustard plants for salt tolerance.     

Sex ratio and spatial pattern of males and females in the dioecious sandhill shrub, Ceratiola ericoides ericoides (Empetraceae) Michx

Sex ratio, size, age, and spatial pattern were investigated for males and females of the dioecious shrub Ceratiola ericoides ericoides (Empetraceae) Michx. within seven mapped populations in Georgia and South Carolina, USA. Among the sites studied, two are regularly burned and one site long-unburned. Age was estimated from node counts of individual shrubs. Only one (fire-suppressed) population showed a female-biased sex ratio, while all others did not differ significantly from 1:1. Mean age estimates did not differ between sexes at any site nor did mean shrub canopy diameter. Bivariate Ripley’s K analysis with a null hypothesis of random labeling was used to investigate whether any of the mapped populations exhibited spatial segregation of the sexes (SSS). No population showed strong evidence of SSS. Rather all sites but one showed males and females to be associated (though not significantly) at a scale of 1–10 m. At a scale of 10–35 m male and female shrubs were located randomly with respect to one another at all sites.     

Influence of irrigation rhythm and water supply on growth,water status and yield of egg-plant (Solanum melongena L.)

Summary The responses of two egg-plant varieties to irrigation rhythm and water supply based on daily potential evapotranspiration were performed on St-Paul silt loam.Results showed that a shorter irrigation rhythm gives the best production in terms of fruit yield (Table 1). Daily irrigation gives a leaf blade with a lower specific weight than the bi-weekly irrigation rhythm during the whole sampling period (Fig. 1).When we compared 4 quantities of water representing 40 to 100% of the potential evapotranspiration, the best results in terms of fruit yield were obtained with the irrigation giving 80% of the potential evapotranspiration (Table 2). But there is no significant difference between the specific leaf weights (Fig. 2).When we observed the varietal behaviour we found that the cultivar Violette de Barbentane gives a production 20% higher than that of Ronde de Valence. But the water potential and transpiration resistance of leaves are not very different for the two varieties (Figs. 5 and 6).The results about the influence of varieties come from one part of the study by M. Pochard and Serieys of the Station of Genetics.     

A comparison of reproductive allocation and reproductive effort between semelparous and iteroparous fucoids (Fucales,Phaeophyta)

Reproductive allocation, the proportion of total dry weight allocated to receptacle tissue and reproductive effort, the proportion of reproductive carbon requirement contributed by receptacle photosynthesis, were measured in two fucoid algal species Fucus serratus and Himanthalia elongata at sites in NE Scotland. Reproductive development takes over ten months in H. elongata, a semelparous (single reproductive event) species, and reproductive allocation at receptacle maturity is over 98%. Following gamete release, the whole thallus dies. In contrast, reproductive development in F. serratus takes four months. Fucus serratus is iteroparous (capable of multiple reproductive events), reproductive allocation is 38.6% for the first reproductive event and 50.5% for the following year's event. In Fucus serratus, the receptacles appear to make a major contribution to their own carbon requirements after the first month of reproductive development. The receptacles of Himanthalia elongata contribute only a small proportion of the receptacle carbon requirements in the first four months of reproductive development, after which the contribution made through receptacle photosynthesis increases.     

Effect of defoliation on leaf growth,sexual expression and reproductive success of Cnidoscolus aconitifolius (Euphorbiaceae)

Several studies on herbivory have shown that plants may have different responses to foliar damage. Most studies have been done using hermaphroditic species, here we present, using a defoliation experiment, different types of response to foliar damage by the monoecious Cnidosculus acontifolius. We evaluated four treatments: natural herbivory (control), and 25%, 50% and 100% artificial defoliation. The results suggest that this species is capable of showing the following compensatory responses: (a) undercompensation in total production of flowers and in fruit set (damage >25%); (b) full compensation in seed weight (all levels of damage); and (c) overcompensation in leaf growth and in the proportion of female flower production (damage 25%). The results show that C. aconitifolius can exhibit a very fast and plastic response depending on the intensity of damage and the time since defoliation. The increase in foliar growth suggests a long-term tolerance response to hervibory, whereas the change in the proportion of female flowers produced does not appear to have a long-term effect due to the decrease of fruit production in all levels of damage. We suggest that the response in foliar growth and the change in sexual expression by C. aconitifolius, are associated with the high resource availability conditions prevalent in the population studied.     

Patterns of growth,reproduction, defense,and herbivory in the dioecious shrub Baccharis halimifolia (Compositae)

Summary Patterns of growth, reproduction, defense (leaf resin) and herbivory were compared between the sexes of the dioecious shrub Baccharis halimifolia (Compositae). Male plants possessed longer shoots and more tender leaves, grew faster, and flowered and senesced earlier than female plants. Levels of leaf nitrogen, water content, and acetone-soluble resin (shown to deter feeding by polyphagous insect herbivores) did not differ between male and female plants. When offered a choice between leaves from male and female plants, adults of two leaf beetles (Chrysomelidae), the monophagous Trirhabda bacharidis and the polyphagous Paria thoracica, both preferred to feed on male leaves. Similarly, the daily fecundity of older females of T. bacharidis was higher when they were fed leaves from male compared to female plants. However, adult survivorship and total fecundity of T. baccharidis did not differ between male and female leaf treatments. We attribute the feeding preference for and slight increase in fecundity on male plants to the tenderness of male leaves. Larvae of the fly Tephritis subpura (Tephritidae) fed exclusively in the sterile receptacle of male flower heads (85% infested), but the phenology was such that pollen production was not adversely affected. Larvae of two other flies Dasineura sp. and Contarinia sp. (Cecidomyiidae) occupied >95% of only female flower heads where they fed among and on the developing seeds. We conclude that foliage-feeding herbivores are unlikely candidates to explain the female-biased sex ratio (59% female) of B. halimifolia plants in the field, and that their preference for male plants is a result of plant characteristics (e.g. rapid growth) that have been selected by some other factor. However, our data on selective floral herbivory in B. halimifolia are in accord with the argument that dioecy reduces the inadvertent loss of flower parts of one sex when herbivores feed on flower parts of the opposite sex.     

Germination potential,growth patterns and reproductive effort of Juncus articulatus and Glyceria australis in temporary shallow wetlands in Australia

Juncus articulatus is an exotic aquatic species occuring over large areas of Mother of Ducks Lagoon, a shallow and temporary wetland in northern New South Wales, Australia. The other dominant perennial species at the site is the native Glyceria australis. Little is known of the ecology of either species or of the potential of J. articulatus to invade wetlands. The germination potential, growth patterns and reproductive effort of J. articulatus and G. australis were compared in response to water regime, grazing and competition. Both species have persistent seed banks from which only a small percentage of seeds germinate in repeated wetting events and from sediment stored dry. J. articulatus germinated in very high numbers from Mother of Ducks Lagoon sediment, but was rare in other lagoons in the region. G. australis germinated in comparatively lower numbers from Mother of Ducks Lagoon sediment and was rare in other lagoon sediments despite being present in the extant vegetation in several lagoons. The introduced J. articulatus has several life cycle characteristics that differ from G. australis, notably a high allocation of biomass to inflorescences under all water regimes. Clipping stimulated above ground vegetative and sexual reproduction for J. articulatus and high numbers of tillers facilitate the sexual reproductive effort. Because J. articulatus is ubiquitous in the region, our results suggest that management of this exotic species should be directed towards minimizing recruitment opportunities and that this could be achieved by increasing variation in water regimes in combination with reduced disturbance from grazing.     

Variation of reproductive effort between parthenogenetic and sexual populations of the scorpion Tityus columbianus

Abstract. Geographic differences in reproductive effort were examined between sexual and parthenogenetic populations of the Colombian scorpion Tityus columbianus , from Iza (sexual) and from Mosquera (unisexual). The sexual females from Iza were significantly larger and had significantly greater relative litter mass (RLM) than the parthenogenetic ones from Mosquera. In both populations litter size increased significantly with female body size.
Litter size and RLM in the sexuals averaged 12.7 and 0.49 respectively, and in the unisexuals 8 and 0.36. We suggest that the higher values are related to greater primary productivity at the Iza site.     

Age, growth, and reproductive biology of the Waigieu seaperch Psammoperca waigiensis (Perciformes: Latidae) around Okinawa Island, Japan

Age, growth, and reproductive biology of the Waigieu seaperch Psammoperca waigiensis were studied using 291 specimens obtained around Okinawa Island, Japan. Otolith opaque zones that formed every year correlated with spawning activity and were thought to be annual rings. Growth of this species was rapid during the first 2 years, reaching 186.2–270.3 mm in standard length (SL). Females (196.6–334.0 mm SL) were larger than males (186.2–288.6 mm SL), caused by differential growth between sexes, which started before 2 years of age. Most of the specimens were 1–11 years old and accounted for 96% in total. Spawning season was estimated to be from April to October by gonadosomatic index (GSI) and histological observation. The smallest mature female and male were 217.0 mm SL (2 years) and 206.0 mm SL (2 years), respectively. After recruitment in rocky areas up to about 200.0 mm SL and 2 years of age, Psammoperca waigiensis were then found to soon mature.     

Flower development and reproductive continuity in MediterraneanRuscus aculeatus L. (Liliaceae)

Summary Ruscus aculeatus is a subandroecious species widespread in Mediterranean environments. Our studies on floral differentiation show that the unisexual flowers pass through a stage in which both androecial and gynoecial primordia are initiated. The hypothesis that dioecy has arisen secondarily from hermaphroditism is discussed. Data on flowering phenology of Mediterranean populations ofR. aculeatus showed it has a long flowering season (about 7 months). This phenology is similar to that of the Mediterranean species originated before the development of the Mediterranean climate. Members of the family Ruscaceae where present in Laurasia during the tropical Tertiary and the present study on flower and fruit morphology and reproductive phenology reveals forR. aculeatus a list of characters generally reported for tropical species. Small and greenish flowers, fleshy fruits, few large seeds, and resprouting capacity, together with long flowering season and continuous availability of ripe fruits, occur all together inR. aculeatus. It constitutes a tropical reproductive syndrome which might have survived the climatic fluctuations of the Quaternary, the establishment of the Mediterranean climate, and the present anthropogenic disturbs.     

Effect of cadmium on the growth and antioxidant enzymes in two varieties of Brassica juncea

Increasing contamination and higher enrichment ratio of non-essential heavy metal cadmium (Cd) induce various toxic responses in plants when accumulated above the threshold level. These effects and growth responses are genotype and Cd level dependent. An experiment was conducted to analyze the effect of Cd toxicity in Brassica juncea [L] Czern and Coss by selecting its two varieties Varuna and RH-30. Cadmium (0, 25, 50 or 100 mg CdCl₂ kg⁻¹ of soil) fed to soil decreased the values of growth characteristics, activity of nitrate reductase and leaf water potential, whereas activities of antioxidant enzymes and proline content increased with the increasing concentration of Cd, observed at 30 and 60 day stages of growth, in both the varieties. Moreover, Cd uptake by the roots was higher in RH-30 than Varuna. Also the activity of antioxidant enzymes and proline accumulation were higher in Varuna with increasing soil level of Cd. Out of the two varieties, Varuna was more tolerant than RH-30 to Cd stress.     

Fecundity,reproductive effort,and pupal size in the profundal midgeChironomus cuccini (Diptera: Chironomidae)

Fecundity ofChironomus cucini increased linearly across a three-fold range of female pupal biomass. Females from a lake producing small pupae (Crystal Lake) had an average of 303 primary follicles, while larger pupae from Trout Lake had an average fecundity of 582. Pupae produced in Crystal Lake from larvae that received experimental food supplements were intermediate in size and fecundity. Pupal biomass explained 76% of the variation in fecundity. No difference in egg size was detected between pupae from the two lakes, and an average value of 2.1 g egg^–1 was used to calculate the total biomass allocated to eggs in each of 79 females dissected. This measure of reproductive effort scaled isometrically with pupal mass. On average, females allocated 48% of their biomass to eggs and 52% to somatic tissues. A comparison of average male biomass to female somatic biomass across 12 populations indicated that the biomass difference between the sexes is a biased predictor of fecundity.