Evolutionary ecology of egg size and number in a seed beetle: genetic trade-off differs between environments

Abstract In many organisms, large offspring have improved fitness over small offspring, and thus their size is under strong selection. However, due to a trade-off between offspring size and number, females producing larger offspring necessarily must produce fewer unless the total amount of reproductive effort is unlimited. Because differential gene expression among environments may affect genetic covariances among traits, it is important to consider environmental effects on the genetic relationships among traits. We compared the genetic relationships among egg size, lifetime fecundity, and female adult body mass (a trait linked to reproductive effort) in the seed beetle, Stator limbatus , between two environments (host-plant species Acacia greggii and Cercidium floridum ). Genetic correlations among these traits were estimated through half-sib analysis, followed with artificial selection on egg size to observe the correlated responses of lifetime fecundity and female body mass. We found that the magnitude of the genetic trade-off between egg size and lifetime fecundity differed between environments–a strong trade-off was estimated when females laid eggs on C. floridum seeds, yet this trade-off was weak when females laid eggs on A. greggii seeds. Also differing between environments was the genetic correlation between egg size and female body mass–these traits were positively genetically correlated for egg size on A. greggii seeds, yet uncorrelated on C. floridum seeds. On A. greggii seeds, the evolution of egg size and traits linked to reproductive effort (such as female body mass) are not independent from each other as commonly assumed in life-history theory.     

Bet-hedging as an evolutionary game: the trade-off between egg size and number

Bet-hedging theory addresses how individuals should optimize fitness in varying and unpredictable environments by sacrificing mean fitness to decrease variation in fitness. So far, three main bet-hedging strategies have been described: conservative bet-hedging (play it safe), diversified bet-hedging (don’t put all eggs in one basket) and adaptive coin flipping (choose a strategy at random from a fixed distribution). Within this context, we analyse the trade-off between many small eggs (or seeds) and few large, given an unpredictable environment. Our model is an extension of previous models and allows for any combination of the bet-hedging strategies mentioned above. In our individual-based model (accounting for both ecological and evolutionary forces), the optimal bet-hedging strategy is a combination of conservative and diversified bet-hedging and adaptive coin flipping, which means a variation in egg size both within clutches and between years. Hence, we show how phenotypic variation within a population, often assumed to be due to non-adaptive variation, instead can be the result of females having this mixed strategy. Our results provide a new perspective on bet-hedging and stress the importance of extreme events in life history evolution.     

Assessment of a rapid method,using soil cores,for estimating the amount and distribution of crop roots in the field

Summary A study was made of the relationship between the number of roots (N_r) observed on unit area of the freshly exposed, horizontal faces of soil cores, and the amounts of roots (per unit volume) present in the same cores. Soil cores, 7 cm diameter, were extracted to depths of 1 m from cereal crops in 1976 at three field sites located on clay soils. Sampling was either at the start of stem elongation, or at anthesis. Estimates of root length per unit soil volume (L) were derived from N_r by assuming random orientation of roots in the soil.Values of L were found to be highly correlated with the measured lengths of both the main roots (root axes) and the total roots (axes and laterals) washed from the soil at a given growth stage, for each of the soils. On average, L was 3.3 times the length of root axes washed from the soil, and was 0.42 times the length of total roots, but there was appreciable variation between different growth stages and field sites. Possible factors giving rise to differences between L and the measured lengths of roots are discussed. Estimates of root length from observation of soil cores may nonetheless provide a suitable basis for rapidly comparing therelative distribution of roots down the soil profile under field conditions.     

Quantifying plant biomass and seed production in camelina (Camelina sativa (L.) Crantz) across a large range of plant densities: Modelling approaches

To date, there has been little agreement on supporting the hypothesis that how some key vegetative traits of camelina (Camelina sativa (L.) Crantz var. ‘Soheil’) are dependent on plant biomass. Therefore, the main aim of this investigation was to quantify the relationship between the size of camelina plants and seed production across a broad-range of plant densities through modelling approaches. To make a wide range of plant densities, a fan design was used in eight replicates in an experimental field at Sari Agricultural Sciences and Natural Resources University, Iran. To quantify the relation between plant density and other plant traits, a regression analysis was carried out and the coefficient of determination (R²) was considered to evaluate the goodness of fit model. A power model (y = ax^b) could describe well the relationship between plant density (ranged 113–2905 plants m⁻²) and plant biomass, seed production, number of seeds per plant, stem diameter, and siliques number, with the coefficient of determination (R²) values of 0.85, 0.87, 0.65, 0.64, and 0.90, respectively. The harvest indexes were 13.8%–26.9%, depending on plant density. Seed production per plant was positively correlated to the siliques number (r = 0.85), the branch number (r = 0.80), and the seed number (r = 0.99) which could be key components of camelina seed production per plant. Furthermore, no significant correlation was found among plant height, thousand-seed weight, and harvest index with seed production per plant. In conclusion, plant biomass could be considered an important trait to predict plant growth models of camelina. Also, a lower plant density of camelina can be compensated by a greater number of siliques, branches and seeds per plant.     

Niche breadth, competitive strength and range size of tree species: a trade-off based framework to understand species distribution

Understanding the mechanisms causing latitudinal gradients in species richness and species range size is a central issue in ecology, particularly in the current context of global climate change. Different hypotheses have been put forward to explain these patterns, emphasizing climatic variability, energy availability and competition. Here we show, using a comparative analysis controlling for phylogeny on 234 temperate/boreal tree species, that these hypotheses can be included into a single framework in an attempt to explain latitudinal gradients in species range size. We find that species tend to have larger ranges when (i) closer to the poles, (ii) successionally seral, (iii) having small and light seeds, and (iv) having short generations. The patterns can simply be explained by energy constraints associated with different life-history strategies. Overall, these findings shed a new light on our understanding of species distribution and biodiversity patterns, bringing new insights into underlying large-scale evolutionary processes.     

Is there a trade-off between egg weight and clutch size in wild Lesser Snow Geese (Anser c. caerulescens)?

Previous studies of arctic nesting geese suggest that laying is limited by the size of a female's body reserves and that larger eggs contain more nutrients. These observations imply a life-history trade-off between egg size and clutch size which may give rise to a negative genetic correlation between the two characters. We estimated the genetic correlation between egg weight and clutch size using measurements from mothers and their daughters in a wild population of Lesser Snow Geese Anser caerulescens caerulescens. Between 65 and 80 % of the variance in egg weight is attributable to differences between individuals, and heritability of egg weight is about 60 %. In contrast, 10–20 % of the variance in clutch size is attributable to differences between individuals, and heritability of clutch size is about 15 %. The genetic correlation coefficient between egg weight and clutch size ranges from 0.09 to 0.32 and does not differ significantly from zero. We discuss the possible reasons for the lack of the expected negative genetic correlation.     

The relationship between body weight and natural mortality in juvenile and adult fish: a comparison of natural ecosystems and aquaculture

The relationship between body weight and natural mortality in juvenile and adult fish was analysed for different aquatic ecosystems: lakes, rivers, the ocean, and pond, cage and tank aquaculture systems. Mortality was modelled as a power function of weight, and the parameters b (exponent) and M_u (mortality at the unit weight of 1 g) estimated for fish in the six ecosystems, as well as within selected populations, species and families. At the ecosystem level, no significant differences in parameters were found between lakes, rivers and the ocean and a joint mortality-weight relationship for all natural ecosystems was estimated with parameters b=?0.288 (90% CL[?0.315, ?0.261]) and M_u=3.00 (90% CL[2.70, 3.30]) year^?1. Among the culture systems, mortality-weight relationships in ponds and cages were not significantly different and a joint relationship was estimated. The weight exponents of mortality in ponds/cages and tanks were very similar at about b=?0.43, and significantly more negative than in natural ecosystems. Mortalities at unit weight were significantly lower in tanks (0.91 year^?1) than in ponds/cages (2.24 year^?1), and both were significantly lower than in natural ecosystems. No systematic differences were found between the mortality-weight relationships determined for individual populations, species or families, and fish in the respective ecosystems. It is hypothesized that aquaculture mortality-weight relationships indicate the allometric scaling of non-predation mortality, which is therefore more strongly size dependent than predation mortality. If non predation mortality in natural ecosystems shows a similar scaling with body weight, then the allometric exponent of predation mortality must be less negative than that observed for total natural mortality. Implications of the established mortality-weight relationships for aquaculture and culture-based fisheries are discussed.     

Domestication and crop physiology: roots of green-revolution wheat

BACKGROUND AND AIMS: Most plant scientists, in contrast to animal scientists, study only half the organism, namely above-ground stems, leaves, flowers and fruits, and neglect below-ground roots. Yet all acknowledge roots are important for anchorage, water and nutrient uptake, and presumably components of yield. This paper investigates the relationship between domestication, and the root systems of landraces, and the parents of early, mid- and late green-revolution bread wheat cultivars. It compares the root system of bread wheat and 'Veery'-type wheat containing the 1RS translocation from rye. METHODS: Wheat germplasm was grown in large pots in sand culture in replicated experiments. This allowed roots to be washed free to study root characters. KEY RESULTS: The three bread wheat parents of early green-revolution wheats have root biomass less than two-thirds the mean of some landrace wheats. Crossing early green-revolution wheat to an F(2) of 'Norin 10' and 'Brevor', further reduced root biomass in mid-generation semi-dwarf and dwarf wheats. Later-generation semi-dwarf wheats show genetic variation for root biomass, but some exhibit further reduction in root size. This is so for some California and UK wheats. The wheat-rye translocation in 'Kavkaz' for the short arm of chromosome 1 (1RS) increased root biomass and branching in cultivars that contained it. CONCLUSIONS: Root size of modern cultivars is small compared with that of landraces. Their root system may be too small for optimum uptake of water and nutrients and maximum grain yield. Optimum root size for grain yield has not been investigated in wheat or most crop plants. Use of 1RS and similar alien translocations may increase root biomass and grain yield significantly in irrigated and rain-fed conditions. Root characters may be integrated into components of yield analysis in wheat. Plant breeders may need to select directly for root characters.     

Wheat as a dual crop for biorefining: Straw quality parameters and their interactions with nitrogen supply in modern elite cultivars

Agricultural residues, such as straw, offer an opportunity to produce biofuels and chemicals in biorefineries without compromising food production. The ideal “dual‐purpose cultivar” would have high yield of grain and straw. In addition, the straw should be easy to process in a biorefinery: It should have good degradability, high concentration of carbohydrates, and low concentration of ash. Nitrogen (N) is an essential nutrient important for plant growth, crop yield and grain quality. However, N production and application comes with a high cost and high environmental footprint. The N application should consequently be based on an economical optimum. Limited knowledge exists on how N application affects the potential of straw for biorefining, for example, straw yield and quality. This study, conducted over three cropping seasons, investigated the effect of N supply on the biorefining potential and included 14 wheat cultivars and one triticale cultivar. The N supply directly affected the yield of straw and grain. In addition, the protein concentration in grain and straw increased, but the composition of the straw with respect to carbohydrates and lignin was largely unaffected by N supply. The only significant change was a lower silicon concentration at increasing N application rate, which could be beneficial for lignin valorization in biorefineries. Likely due to the negligible changes in cell wall composition, the effect of N application rate on straw degradability was not significant. N application should therefore primarily be optimized with respect to grain quality and overall yield of grain and straw. Differences between cultivars were also minor with respect to their performance in a biorefinery process. From a breeding and agronomic perspective, focus should therefore be put on maximizing the biomass output from the field, that is, selecting the cultivar with highest grain and straw yield and optimizing the application of fertilizer to get optimum N use efficiency.     

Abundance and habitat segregation in Mediterranean grassland species: the importance of seed weight

Abstract. We analysed the relationship between seed traits (weight, shape and dispersal structures) and the abundance and habitat segregation of Mediterranean grassland species. To take into account possible correlations with other plant traits, the study also includes 5 vegetative traits (growth form, plant longevity, clonality, onset of flowering and plant size) of commonly accepted functional importance. Data were recorded for 85 species from dehesa grasslands in central Spain. Species abundance was measured in upper (dry and less productive, high stress) and lower (moist and more productive, low stress) slope zones in the same area. Habitat segregation was estimated using an index based on the relative frequencies of species in upper and lower slope zones. Multiple regression models were fitted using species, as well as phylogenetically independent contrasts, as data points. Annual small‐seeded species without specialised dispersal structures are over‐represented in dehesa grasslands. Abundance was negatively related to seed weight in upper slope zones. None of the recorded plant traits were related to abundance in the lower slope zones. Habitat segregation was mainly related to seed weight, but also to some vegetative traits. Annual, early flowering and small‐seeded species were relatively more abundant in the upper than the lower slope zones. This pattern is independent of phylogeny. Our results suggest that in dry Mediterranean grasslands, abundance of many species is determined by dispersal (production of numerous small seeds) rather than by competitive ability.     

Harvest index: a review of its use in plant breeding and crop physiology

This review charts the use of the concept of harvest index in crop improvement and physiology, concentrating on the literature from the last 20 years. Evidence from abstract journals indicates that the term has been applied most to small grain cereal crops and pulses, in India, Western Europe and the USA, and that it has been less useful for maize and tuber crops. Standard methods of measuring harvest index, the associated problems of measurement and interpretation, and representative values for a range of world species are reviewed. The values for modern varieties of most intensively-cultivated grain crops fall within the range 0.4 to 0.6. Variation between varieties of the same species is illustrated by trends in the harvest indices of old, outclassed and recent varieties of temperate and mediterranean wheat and barley (compared under uniform conditions); this shows a progressive increase throughout the present century, although improvement has been much slower in Australia and Canada than in the UK. In most cases, the improvement in harvest index has been a consequence of increased grain population density coupled with stable individual grain weight. The high heritability of harvest index is explored by examining its (rather weak) response to variation in environmental factors (fertilisation, population density, application of growth regulators) in the absence of severe stress. A fuller perspective is gained by reviewing aspects of the harvest index of rice, maize and tropical pulses. With rice, attention must be paid to the fact that the adhering lemma and palea (not primarily part of economic yield) can make up 20% of grain weight; and there are important interactions among biomass, grain yield and season length. Maize differs from most small grain crops in that harvest index (in N. American varieties) was already high at the start of this century, and increases in yield potential have been largely the consequence of increased biomass production. The harvest index of many pulse species and varieties tends to be low because selection has been for some yield in all seasons. Extension of the harvest index concept to express the partitioning of mineral nutrients as well as dry matter (e.g. the nitrogen harvest index) has provided a range of responses whose implications for production and breeding remain to be explored. It is concluded that even though the principal cereal crops appear to be approaching the upper limit of harvest index, and future yield gains will have to be sought by increased biomass production, there will still be a need for the concept of harvest index as a tool in interpreting crop response to different environments and climatic change.     

Daisyworld inhabited with daisies incorporating a seed size/number trade-off: the mechanism of negative feedback on selection from a standpoint of the competition theory

We reexamined a Daisyworld model from the traditional view of competition theory. Unlike the original model, white and black daisies in our model incorporate a seeding/germination trade-off against bare ground area without assuming the local temperature reward. As a result, the planetary temperature is automatically regulated by two species if the following conditions are met: (i) the species react equally to an environmental condition, but one can alter the environmental condition in the opposite direction to the other. (ii) that one of the two cannot have both a higher maximal growth rate (mu(max)) and lower half-saturation constant (K) than those of the other. In other words, a pair of phenotypes incorporates a trade-off between quality and number of seeds. We found that the homeostatic regulation can also be reconciled with the adaptive evolution of optimal temperature. The results of simulation imply that biotic environmental feedback can also be maintained when the emergence of polymorphisms (black and white daisies) is closely linked to such a trade-off.     

Life history evolution in cichlids 2: directional evolution of the trade-off between egg number and egg size

The negative relationship between offspring number and offspring size provides a classic example of the role of trade-offs in life history theory. However, the evolutionary transitions in egg size and clutch size that have produced this negative relationship are still largely unknown. Since body size may affect both of these traits, it would be helpful to understand how evolutionary changes in body size may have facilitated or constrained shifts in clutch and egg size. By using comparative methods with a database of life histories and a phylogeny of 222 genera of cichlid fishes, we investigated the order of evolutionary transitions in these traits in relation to each other. We found that the ancestral large-bodied cichlids first increased egg size, followed by a decrease in both body size and clutch size resulting in the common current combination of a small-bodied cichlid with a small clutch of large eggs. Furthermore, lineages that deviated from the negative relationship between clutch and egg size underwent different transitions in these traits according to their body size (large bodied genera have moved towards the large clutch/small egg end of the continuum and small bodied genera towards the small clutch/large egg end of the continuum) to reach the negative relationship between clutch size and egg size. Our results show that body size is highly important in shaping the negative relationship between clutch size and egg size.     

A triangular relationship between leaf size and seed size among woody species: allometry, ontogeny, ecology and taxonomy

 A hypothesized relationship between seed weight and leaf size was investigated for 58 diverse British (semi-)woody species. Interspecific variation in leaf size of adult plants corresponded allometrically with interspecific variation in the weight of an infructescence (seed-bearing inflorescence). The relationship between seed size and leaf size of adult plants was triangular. The corners of the triangle were interpreted in terms of ecological strategy. Medium-sized infructescences, small seeds and large leaves were seen among medium-sized, fast-growing, earlier-successional, mostly deciduous shrubs and trees; small infructescences, small seeds and small leaves mostly among low, slow-growing evergreens from stress-prone, proclimax habitats; and large infructescences, large seeds and large leaves among slow-growing, later-successional trees of potential competitive vigour. The hypothesis that the combination of large seeds and small leaves is allometrically unlikely was supported by the data. The roles of ontogeny and taxonomic relatedness in the seed size-leaf size relationship were examined by correlative and taxonomic analyses of seed, plant and leaf size during the unfolding of the life history from seed through two seedling phases to adulthood. Deciduous versus evergreen leaf habit was a source of deviation from the otherwise linear allometric relationships during ontogenetic development, none of which were, individually, confounded significantly with taxonomy. Received: 2 March 1998 / Accepted: 15 October 1998     

Interaction between algae and seed germination in tropical dune slack species: a facilitation process

The importance of algae dune slack stabilisation has been extensively analysed, but little has been done regarding their potentially favourable role in germination and establishment of phanerogams. Dune slack is a seasonal freshwater habitat that depends on water table level fluctuation. Our hypothesis is that in these highly unpredictable systems, algae maintain a certain degree of humidity in the soil and, thus, permit phanerogam germination and establishment. The objective of this study was to compare the germination of four species characteristic of tropical dune systems: Cyperus articulatus L., Fimbristylis cymosa R. Br., Fuirena simplex Vahl and Rhynchospora colorata (L.) Pfeiffer, in different substrates and degrees of humidity. For germination trials, seeds of the four species were placed each on three substrates: in sand (control), filamentous algae on sand and cotton on sand. Two irrigation protocols were followed: continual watering (wet treatment) and weekly watering (dry treatment). Each week during a 21-week period, the number of germinated seeds from each species was recorded. The highest percentage of germination for the four species was observed during wet treatment. C. articulatus had the highest germination rate of the four species in algae (70%), and cotton (51%) on sand. Highest germination for F. simplex, F. cymosa and R. colorata took place in algae, although the percentages were very low. Differences in germination between the three substrates for these species were statistically significant (P<0.05). During dry treatment, C. articulatus was the only species that exhibited a significant germination, principally in algae and cotton. The results suggest that algae maintain substrate humidity and favour germination of the four species. This could be important in sand dune slacks where marked fluctuations in temperature and humidity occur.     

The relationship between species richness and standing crop in wetlands: the importance of scale

One of the few important empirical generalizations regarding herbaceous plant systems has been the demonstration that species richness is related to standing crop with maximum richness occurring at moderate levels of standing crop. This relationship is normally demonstrated by comparing among vegetation types (i.e., vegetation with different dominants). We undertook this study to test whether the species richness-standing crop relationship was evident at a finer-grained level of organization, the within vegetation type level. Fifteen wetland sites were sampled in eastern Canada and species richness and standing crop determined in each of 224 0.25 m² quadrats. Each site was relatively homogeneous in terms of the dominant species present and were therefore categorized as single vegetation types. However, as a group, the sites comprised a wide range of vegetation types.A second order polynomial regression indicated a significant bitonic relationship between species richness and standing crop at the among-vegetation types scale, that is, when all 15 sites were combined. At the within-vegetation type level, however, no significant relationships were observed (p>0.05). The results indicate that the model of species richness proposed by Grime has predictive power at a coarse-grained level of organization, among vegetation types, but does not survive the transition to a finer-grained level of organization, the within vegetation type level. Therefore, the higher level processes which structure species richness patterns among vegetation types are not the same processes which determine richness patterns within a vegetation type.     

The dynamics of three shrub species in a fire-prone temperate savanna: the interplay between the seed bank, seed rain and fire regime

A model was developed to assess how the seed rain and fire regime affect seed bank dynamics and seedling establishment of three native shrub species (Acanthostyles buniifolius, Baccharis pingraea and Baccharis dracunculifolia) with different regeneration strategies, in temperate South American savanna. Seed bank and seed rain were quantified for each species under different fire regimes, and their relative roles in regeneration were evaluated. All species had short-term persistent seed banks and high annual variability in seed production. A high proportion of seeds deposited in the seed rain produced seedlings after fire; few entered the soil seed bank. Fire killed a high proportion of the seeds in the soil seed bank. Seedlings derived from the seed rain had a higher probability of surviving for 2 years than seedlings emerging from the soil seed bank. In the absence of fire, establishment depended on germination both from the seed rain and the soil seed bank, whereas with annual fire, establishment was primarily dependent on germination of seeds arriving in the annual seed rain, regardless of species’ regeneration strategies. These results help to explain changes in the vegetation of South American temperate savannas as a result of changes in fire regime and grazing management during the last 50 years. By revealing the crucial roles of the soil seed bank and seed rain in regeneration, this study provides vital information for the development of appropriate management practices to control populations of shrub species with different regeneration strategies in South American temperate savannas.