Showing papers in "Biological Reviews in 1994"

Social learning in animals : categories and mechanisms

Abstract: There has been relatively little research on the psychological mechanisms of social learning. This may be due, in part, to the practice of distinguishing categories of social learning in relation to ill-defined mechanisms (Davis, 1973; Galef, 1988). This practice both makes it difficult to identify empirically examples of different types of social learning, and gives the false impression that the mechanisms responsible for social learning are clearly understood. It has been proposed that social learning phenomena be subsumed within the categorization scheme currently used by investigators of asocial learning. This scheme distinguishes categories of learning according to observable conditions, namely, the type of experience that gives rise to a change in an animal (single stimulus vs. stimulus-stimulus relationship vs. response-reinforcer relationship), and the type of behaviour in which this change is detected (response evocation vs. learnability) (Rescorla, 1988). Specifically, three alignments have been proposed: (i) stimulus enhancement with single stimulus learning, (ii) observational conditioning with stimulus-stimulus learning, or Pavlovian conditioning, and (iii) observational learning with response-reinforcer learning, or instrumental conditioning. If, as the proposed alignments suggest, the conditions of social and asocial learning are the same, there is some reason to believe that the mechanisms underlying the two sets of phenomena are also the same. This is so if one makes the relatively uncontroversial assumption that phenomena which occur under similar conditions tend to be controlled by similar mechanisms. However, the proposed alignments are intended to be a set of hypotheses, rather than conclusions, about the mechanisms of social learning; as a basis for further research in which animal learning theory is applied to social learning. A concerted attempt to apply animal learning theory to social learning, to find out whether the same mechanisms are responsible for social and asocial learning, could lead both to refinements of the general theory, and to a better understanding of the mechanisms of social learning. There are precedents for these positive developments in research applying animal learning theory to food aversion learning (e.g. Domjan, 1983; Rozin & Schull, 1988) and imprinting (e.g. Bolhuis, de Vox & Kruit, 1990; Hollis, ten Cate & Bateson, 1991). Like social learning, these phenomena almost certainly play distinctive roles in the antogeny of adaptive behaviour, and they are customarily regarded as 'special kinds' of learning (Shettleworth, 1993).(ABSTRACT TRUNCATED AT 400 WORDS)

Intraspecific variation in egg size and egg composition in birds: effects on offspring fitness

Abstract: 1. There is little unequivocal evidence to date in support of a positive relationship between egg size and offspring fitness in birds. Although 40 studies (of 34 species) have considered the effect of variation in egg size on chick growth and/or survival up to fledgling only 12 studies have controlled for other characters potentially correlated both with egg size and offspring fitness. Of these only two have reported a significant residual effect of egg size on chick growth (in the roseate tern and European blackbird) and three a residual effect on chick survival (all in seabirds: common tern, lesser black-backed gull and kittiwake). 2. More consistent evidence exists, though from fewer studies, for a positive relationship between egg size and offspring fitness early in the chick-rearing period; chick growth and chick survival being dependent on egg size in 8 of 10 studies and 4 of 5 studies respectively. It is suggested that the most important effect of variation in egg size might be in determining the probability of offspring survival in the first few days after hatching. 3. Egg size explains on average 66% of the variation in chick mass at hatching (n = 35 studies) but only 30% of the variation in chick body size (n = 18). When effects of hatching body size are controlled for chick mass remains significantly correlated with egg size, though the reverse is not true. This supports the hypothesis that large eggs give rise to heavier chicks at hatching, i.e., chicks with more nutrient (yolk) reserves, rather than structurally larger chicks. 4. Egg composition increased isometrically with increasing egg size in about half the studies so far reported (n equals approximately 20). However, in seabirds, and some passerines, larger eggs contain disproportionately more albumen, whilst in some waterfowl percentage yolk content increases with increasing egg size. Changes in albumen content largely reflect variation in the water content of eggs, but changes in yolk content involve variation in lipid content, and therefore in egg 'quality.' The adaptive significance of variation in egg composition is considered; females may adjust egg composition facultatively to maximise the benefits to their offspring of increased reproductive investment. 5. Considerations for future research are discussed with particular emphasis on experimental studies and the application of new techniques.

Differential fitness returns in relation to spatial position in groups

Abstract: 1. A general overview of position-related fitness differences in group-living animals is given for the first time. 2. Differential fitness returns in groups are often related to competition for resources among group members due to resource limitation. If resources are very scarce, then competition may finally lead to the disbandment of groups. The encounter-dilution effect predicts that grouping confers a benefit if the conspicuousness of large groups does not outweigh the dilution effect and each predator or parasite attacks only one (or a few) prey per encounter (Uetz & Hieber, 1993). While the encounter-dilution effect limits the number of predators and parasites a group is faced with at a given time, marginal predation and communal defence are probably the most important factors for position-related differences in predation risk. 3. Limitation of resources increases competition between groups members and should widen the gap between fitness returns of dominant and subordinate group members. In some group-living animals such as schooling fish, however, predator pressure has apparently selected for uniformity in morphology and behaviour which presumably restricts the potential for resource competition (Magurran & Seghers, 1991). 4. A general problem of assessing individual fitness returns in that different short-term strategies may achieve the same long term goals (Magurran, 1993). The use of feeding rates and predation risks as currencies for fitness returns over short time periods is therefore questionable and can only be a first step. So far, hardly any information is available on long-term effects of differences in positioning behaviour. 5. Food availability and food intake rates tend to be higher in edge positions, whereas energy expenditure does not differ significantly with group position in non-roosting mobile groups. This suggests that edge positions achieve a higher net-energy pay-off and should therefore be preferred by individuals with low energy reserves. 6. Differential predation risks are well documented for stationary and colony-breeding species, but there is a lack of data for mobile, non-breeding species. The reason for the latter is that predation is rarely observed (in the field) and often difficult to relate to group position in fast-moving species such as flocks of birds. Further studies, especially in the field, are needed. 7. Vigilance rates and attack rates do not necessarily give a true reflection of mortality risk. Vigilance is not exclusively related to predation and can be confounded by competition effects and hunger. Attack rates are difficult to translate into per capita risks if hardly any mortality is observed and therefore little information is available about the ratio of successful attacks/total attacks. 8. Most of the literature up to date consists of descriptive studies. There is a need for experimental work that investigate (a) the influence of predator attack mode on differential predation risks; (b) the influence of internal states (such as hunger) on the tradeoff between predation risk and foraging behaviour and its consequences for the positioning behaviour in groups; (c) the 'egalitarian' nature of groups with respect to group position. Do all individuals continuously rotate positions? Or do dominant individuals monopolize certain positions?, and (d) that separate the effects of nutritional state, vigilance and food availability on feeding rates in different group positions. 9. Reproductive success is often related to group position in breeding colonies and lekking animal species. Females probably prefer the safer territories in the group centre, but more empirical data are required to test this hypothesis. 10. Position-related differences in parasitism rate between group members are well documented with per capita rate of parasitism being lower in the centre of groups than at the periphery.(ABSTRACT TRUNCATED AT 400 WORDS)

Heterospory: the most iterative key innovation in the evolutionary history of the plant kingdom

Abstract: Summary 1In aggregate, past discussions of heterospory and its role in the alternation of generations are riddled with ambiguities that reflect overlap of terms and concepts. Heterospory sensu lato can be analyzed more effectively if it is fragmented into a series of more readily defined evolutionary innovations: heterospory sensu stricto (bimodality of spore size), dioicy, heterosporangy, endospory, monomegaspory, endomegasporangy, integumentation, lagenostomy, in situ pollination, in situ fertilization, pollen tube formation, and siphonogamy (Tables 1, 2, Figs 1, 13). Current evidence suggests that the last five characters are confined to the seed-plants. 2The fossil record documents repeated evolution of heterosporous lineages from anisomorphic homosporous ancestors. However, interpretation is hindered by disarticulation of fossil sporophytes, the difficulty of relating conspecific but physically independent sporophyte and gametophyte generations in free-sporing pteridophytes, the inability to directly observe ontogeny, and the rarity of preservation of transient and/or microscopic reproductive phenomena such as syngamy and siphonogamy. Unfortunately, the rarely preserved phenomena are often of far greater biological significance than corresponding readily preserved phenomena (e.g. heterospory versus dioicy, heterosporangy versus endospory). 3In most fossils gametophyte gender can only be inferred by extrapolation from the morphology of the sporophyte and especially of the spores. This is readily achieved for species possessing high-level heterospory, when the two spore genders have diverged greatly in size, morphology, ultrastructure and developmental behaviour. However, the earliest stages in the evolution of heterospory, which are most likely to be elucidated in the early fossil record of land-plants, also show least sporogenetic divergence. It is particularly difficult to distinguish large microspores and small megaspores from the large isospores of some contemporaneous homosporous species (Figs 3–6 a, g). Heterospory is best identified in fossils by quantitative analysis of intrasporangial spore populations. 4The spatial scale of the differential expression of megaspores and microspores varies from co-occurrence in a single sporangium (anisospory) to different sporophytes (dioecy) (Figs 6–8). Studies of the relative positions of the two spore morphs on the sporophyte, and of developmentally anomalous terata (Fig. 9), demonstrate that gender is expressed epigenetically in both the sporophyte and gametophyte. Hormonal control operates via nutrient clines, with nutrient-rich microenvironments favouring femaleness; megaspores and microspores compete for sporophytic resources. External environments can also influence gender, particularly in free-living exosporic gametophytes. 5The evolution of heterospory was highly iterative. The number of origins is best assessed via cladograms, but no current phylogeny includes sufficient relevant tracheophyte species. Also, several extant heterosporous species differ greatly from their closest relatives due to high degrees of ecological specialization and/or saltational evolution; extensive molecular data will be needed to ascertain their correct phylogenetic position. Current evidence suggests a minimum of 11 origins of heterospory, in the Zosterophyllopsida (1: Upper Devonian), Lycopsida (1: Upper Devonian), Sphenopsida (?2: Lower Carboniferous), Pteropsida (?4: Upper Cretaceous/Palaeogene) and Progymnospermopsida (?3: Upper Devonian/Carboniferous). The arguably monophyletic Gymnospermopsida probably inherited heterospory from their progymnospermopsid ancestor (Table 3, Figs 11–13). No origin of heterospory coincides with the origin of (and thus delimits) any taxonomic class of tracheophytes. The actual number of origins of heterospory is probably appreciably higher, exceeding that of any other key evolutionary innovation in land-plants and offering an unusually good opportunity to infer evolutionary process from pattern. 6Heterospory reflects the convergent attainment of similar modes of reproduction in phylogenetically disparate lineages. Nature repeated this experiment many times, whereas true phylogenetic synapomorphies evolve only once and require a unique causal explanation. Cladograms also offer the best means of determining the sequence of acquisition of heterosporic phenomena within lineages, here exemplified using the lycopsids (Fig. 10). Comparison of such sequences among lineages can potentially allow generalizations about underlying evolutionary mechanisms. Current evidence (albeit inadequate) indicates broadly similar sequences of character acquisitions in all lineages, though they differ in detail. Some logical evolutionarily stages were temporarily bypassed. Other lineages appear to have acquired two or more characters during a single saltational evolutionary event. Heterosporic phenomena can also be lost during evolution. Although no complete reversals to homospory have been documented, this could reflect unbreakable developmental canalization of heterospory rather than selective advantage relative to homospory. Several extant species refute widely held assumptions that certain phenomena, notably heterospory and dioicy, are reliably positively correlated. Moreover, some fossils are likely to possess combinations of heterosporic characters that are not found in their extant descendants. Fossil data have played a crucial role in understanding both the number of origins of heterospory and the ensuing patterns of character acquisition. 7Although non-adaptive evolutionary events are likely in at least some lineages, the highly iterative nature of heterospory and similar patterns of character acquisition in different lineages together suggest that its evolution was largely adaptively driven. However, many previous adaptive models of heterosporic evolution confused pattern and process, and paid insufficient attention to the role of the environment as a passive filter of novel morphotypes. Linear gradualistic models were imposed on the data, often intercalating hypothetical intermediates where desired. 8The evolution of heterospory is best understood in terms of inherent antagonism between the sporophytic and gametophytic phases of the life history for control of sex ratio and reproductive timing. Control is achieved directly by the gametophyte, via gametogenesis, and indirectly by the sporophyte, via sporogenesis and the ability to determine to varying degrees the environment in which the gametophyte undergoes sexual reproduction. Increasing levels of heterospory (particularly the acquisition of endospory) compress the heteromorphic life history, as the increasingly dominant sporophyte progressively co-opts the sex determination role of the gametophyte. The resulting life history is more holistic, effectively streamlining evolution by offering only a single target for selection. 9However, by wresting control of sex ratios from the gametophyte, the ability of the sporophyte to respond rapidly to environmental changes decreases. This competitive weakness is greatest for heterosporous species possessing exosporic but obligately unisexual gametophytes (epitomized by the pteropsid Platyzoma*). It can be alleviated in endosporic species by occupying favourable environments (e.g. the aquatic Salviniales and Marsileales), switching to an apomictic mode of reproduction (thereby incurring inbreeding depression; e.g. many selaginellaleans), or acquiring more complex pollination biologies (thereby by-passing the environment as a selective filter: the seed-plants). 10Lineages differ greatly in the maximum number of heterosporic characters that were acquired by their most derived constituent species. Several Devono-Carboniferous lineages reached the level of reducing numbers of functional megaspores to one per sporangium (Figs 7 e, f, 8, 13), but only the putatively monophyletic gymnospermopsids broke through this apparent barrier to acquire the increasingly complex pollination biology that characterizes modern seed-plants. 11Many theories have been proposed to explain the remarkable success (both in terms of species diversity and ecological dominance) of seed-plants. The majority focus on characters that are absent from the earliest seed-plants (the Devono-Carboniferous lyginopterid pteridospermaleans), which were no more reproductively sophisticated than other penecontemporaneous lineages possessing advanced heterospory (particularly the most derived lycopsids, equisetaleans and progymnospermopsids). Reliable pollination was a key reproductive breakthrough, though the sophisticated economic-vegetative characters inherited by the earliest seed-plants from their putative progymnospermopsid ancestors were probably equally important in ensuring their success in water-limited habitats. 12With the exception of some ecologically specialized pteropsids, known heterosporous lineages originated during a relatively short period in the Upper Devonian and Carboniferous (Fig. 11). They exploited a window of opportunity that existed before niches became too finely partitioned and saturated with seed-plant species. This non-uniformitarian ecology renders negligible the probability of new heterosporous lineages becoming established today, even though ‘hopeful monsters’ possessing ‘incipient heterospory’ are probably constantly being generated from homosporous parents.

Cost of growth in cells and organisms: general rules and comparative aspects

Abstract: In a crude fashion it can be said that metabolizable energy (M) is partitioned into metabolic work, paid for by 'oxidations' (R), and 'assimilation', i.e. production (P), so that M = R+P. However, a fraction of R is required to meet the expenses of production and if these expenses represent, Joule for Joule, a constant proportion of the amount produced, then Rt = Rm+cP, where Rt = total metabolic expenditures, Rm = metabolic expenditures for maintaining the non-producing organism, and cP = Rp = metabolic expenditures connected with the processes of production. The partitioning of metabolizable energy into R and P as well as into Rm and Rp may vary depending on the phylogeny and life-history of the species concerned and on ecological circumstances. Thus selection is expected to act on both ratios, R/P and Rm/Rp. By comparing the ratios P/(P+Rp) (the apparent efficiency of production) and Rp/P (the apparent metabolic cost of production) in different types of organisms, one finds that a value of P/(P+Rp) = 0.75, equal to 75% efficiency, 10 mgdbm/mmol ATP, and 16 mumolO2/mg dbm (when I mg identical to 22 J), can be used as a 'consensus value' for the average efficiency, or cost, of the transformation of metabolizable energy into production in a wide range of organisms, from bacteria to mammals. This value corresponds to about three times the theoretical cost of synthesizing the same amount of tissue on the basis of known biochemical principles. The reasons why the empirical costs of production are higher than the theoretical costs of synthesis by what appears to be a common factor may be quite different in bacteria, small ectothermic and large endothermic organisms. Deviations from the consensus value may be due to differences in energy density of the nutrients assimilated and the tissues synthesized. Further complications arise because of interactions between P, Rp, and Rm. In microorganisms the existence of a constant and a variable component of maintenance metabolism has been postulated, the latter decreasing with increasing rate of production. In small ectothermic metazoans, on the other hand, the nonlinear relationship between growth metabolism and growth rate has led to the speculation that above a critical value of Pg certain energy consuming functions of maintenance are suppressed and the energy thus gained used for fuelling growth processes. There is some evidence that, at least in ectothermic metazoans, the apparent cost of growth decreases with the rate of growth, reaching a low plateau of about 10 mumolO2/mgdbm at growth rates exceeding about 8 mgdbm/g/h.(ABSTRACT TRUNCATED AT 400 WORDS)

The fundamental theorem of natural selection.

Abstract: The fundamental theorem of natural selection was a valiant attempt by R. A. Fisher to fit a quart into a pint pot The present author reviewing Population Genetics by Ewens, 1969a (Edwards, 1969). Many authors have maintained that the theorem holds only under very special conditions, while only a few (e.g. Edwards, 1967) have thought that Fisher may have been correct – if only we could understand what he meant! It will be shown here that this latter view is correct Price, 1972. My own reason for not including a planned chapter on the fundamental theorem is… [that] in spite of many efforts I am still not satisfied that I am able to provide an account that does the theorem justice The present author in the Epilogue to Foundations of Mathematical Genetics (Edwards, 1977).

Biological assessment of aquatic pollution: a review, with emphasis on plants as biomonitors*

Abstract: In a number of disciplines including ecology, ecotoxicology, water quality management, water resource management, fishery biology etc., there is significant interest in the testing of new materials, environmental samples (of water or sediments) and specific sites, in terms of their effects on biota. In the first instance, we consider various sources of aquatic pollution, sources typically associated with developed areas of the world. Historically, much water quality assessment has been performed by researchers with a background in chemistry or engineering, thus chemical analysis was a dominant form of assessment. However, chemical analyses, particularly of such materials as organochlorines and polyaromatic hydrocarbons can be expensive, and local environmental factors may cause the actual exposure of an organism to be little correlated with chemical concentrations in the surrounding water or sediments. To a large extent toxicity testing has proceeded independently of environmental quality assessment in situ, and the work has been done by different, and differently-trained researchers. Here we attempt to bring together the various forms of biological assessment of aquatic pollution, because in our opinion it is worth developing a coherent framework for the application of this powerful tool. Biotic assessment in its most primitive form involves the simple tracking of mortality in exposed organisms. However, in most natural environments it is extended, chronic exposure to contaminants that has the most wide-ranging and irreversible repercussions--thus measures of sub-lethal impairment are favoured. From an ecological standpoint, it is most valuable to assess ecological effects by direct study of in situ contaminant body burdens and impairment of growth and reproduction compared with 'clean' sites. A distinction is made here between bioindication and biomonitoring, and a case is made for including aquatic macrophytes (angiosperms) in studies of contaminant levels and effects in the biota. It is apparent that there is a concurrent need for laboratory-based testing of new industrial by-products before any are released in the environment, and such studies should aid the investigation of mechanisms and modes of toxicity, but environmental assessment, and tracking of improvements in environmental quality are most effectively achieved by active biomonitoring experiments.

Evolutionary-developmental change in the growth architecture of fossil rhizomorphic lycopsids: scenarios constructed on cladistic foundations

Abstract: Summary The Rhizomorphales, the most derived portion of the lycopsid (clubmoss) Glade, is now represented only by the diminutive genus Isoetes. However, during their Late Palaeozoic acme the rhizomorphic lycopsids exhibited a wide range of architectures and body sizes, from recumbent pseudoherbs to trees 40 m high. All possessed the rhizomorphic syndrome: a centralized rootstock and secondary thickening, reflecting an inescapable developmental constraint of bipolar determinate growth. These features in turn allowed acquisition of the tree habit by the lycopsids, independently of the physiologically and ontogenetically distinct lignophyte Glade that includes the seed-plants. Differences among lycopsid genera in the number and size of four major growth modules – rhizomorph, stem, lateral branches (two positionally distinct submodules), and isotomous crown branches – resulted from differences in the relative size, number and equality of dichotomies of the apical meristems. A detailed experimental cladistic analysis of the best known fossil rhizomorphic lycopsids demonstrates extensive iteration among several distinct growth architectures characterizing ten genera. Scenarios can be constructed for the type of morphogenetic transitions necessary to (1) derive one genus from a putative ancestor or (2) explain the relationship of two genera relative to a putative outgroup. The scenarios are best formulated within a synthesis of terminology devised primarily by zoologists to describe via size–shape trajectories various modes of evolutionary–developmental change: heterotopy, heterochrony sensu lato, and allometric modifications. Many examples of these phenomena are evident among the rhizomorphic lycopsids, and can be explored by reconstructing hypothetical ancestors occupying interior nodes of the cladogram. Iterative origination of the small-bodied genera from tree-sized ancestors is inferred, by various forms of paedomorphosis and decreases in the number of developmental stages that are sufficiently profound to locally perturb perceived phylogenetic relationships. This study highlights several problems of cladistic analysis in general and of fossils in particular, especially the significance for character polarization of both perceived primitiveness and large phenetic gaps. Many phylogenetic studies of several extant angiosperm clades also imply frequent architectural transitions, but few suggest repeated origins of non-trees from trees. A simple model for control of development focuses on D-genes: switches that control morphogen production. This perspective emphasizes the importance of a series of inter-related hierarchies reflecting ontogenetic time, size, burden and complexity within species, and phylogeny among species. The increasingly evident simplicity and common origin of D-gene control in all living organisms is used to formulate a neoGoldschmidtian paradigm of instantaneous, non-adaptive, saltational speciation via teratological ‘hopeful monsters’ that escape the constraints of developmental canalization. The scenario does not require large-scale mutations or high levels of fitness in the mutants - merely temporary release from competitive selection by establishing the new lineage in a vacant niche. The model is more appropriate to higher plants than to the more developmentally complex and constrained higher animals, and more appropriate to fossil plants than to their more ecologically complex and developmentally constrained living descendants. Future progress in understanding transitions in plant form requires reciprocal illumination between such scenarios and empirical observations of gene expression analyzed in a cladistic context.

Alternation of generations in land plants: new phylogenetic and palaeobotanical evidence

Abstract: Summary Current ideas on the evolution of alternation of generations in land plants are reviewed in the context of important recent advances in plant systematics and the discovery of remarkable new palaeobotanical evidence on early embryophyte life cycles. An overview of relationships in major groups of green plants is presented together with a brief review of the early fossil record as a prelude to discussing hypotheses of life cycle evolution. Recent discoveries of life cycles in the early fossil record are described and assessed. The newly discovered gametophyte and sporophyte associations are based on exceptionally well-preserved material from the Rhynie Chert, Scotland (Middle Devonian: 380–408 Myr) and compression fossils from other Devonian localities. These data document diplobiontic life cycles in plants at the ‘protracheophyte’ and early tracheophyte level of organization. Furthermore, the early fossils have a more or less isomorphic alternation of generations, a striking departure from life cycles in extant embryophytes. This unexpected similarity between gametophyte and sporophyte calls for a cautious approach in identifying ploidy level in early groups. Viewed in a systematic context, the neontological and palaeontological data contribute towards the formulation of a coherent hypothesis of life cycle evolution in major, early embryophyte groups. Evidence from extant groups strongly supports a single direct origin of the diplobiontic life cycles of land plants from haploid, haplobiontic life cycles in ancestral ‘charophycean algae’. The interest of the new palaeobotanical data lies in its relevance to life cycle evolution at the restricted level of vascular plants rather than at the more general level of embryophytes (vascular plants plus ‘bryophytes’). The occurrence of morphologically complex, axial gametophytes in early vascular plants is consistent with the moss sister-group proposed in some cladistic analyses. Similarities of moss gametophytes to fossils in the vascular plant stem-group are discussed, and it is argued that the late appearance of mosses in the macrofossil record may be due to the problem of recognizing stem-group taxa. The new palaeobotanical evidence conflicts with previous hypotheses based on extant groups that interpret morphological simplicity as the plesiomorphic condition in the gametophytes of vascular plants. These new data indicate that a significant elaboration of both gametophyte and sporophyte occurred early in the tracheophyte lineage, and that the gametophytes of extant ‘pteridophytes’ are highly reduced compared to those of some of the earliest ‘protracheophytes’. Vestiges of this early morphological complexity may remain in the gametophytes of some extant groups such as Lycopodiaceae.

The influence of natural and experimental high o2concentrations on o2-evolving phototrophs

Abstract: CONTENTS

The origin of the land plant sporophyte: an interpolational scenario

Abstract: Summary The origin of terrestrial plants from a charophycean ancestor is assumed as a basis for the consideration of the origin of life histories amongst this group. Charophycean algae are vegetatively gametophytic, thus requiring the interpolation of the multicellular sporophytic stage. The model presented here derives the sporophyte and typical sporangial contents from the zygospore produced from the reproductive structure of a hypothetical charophycean land plant ancestor. The presence of monads, dyads, and various forms of tetrads in the fossil record of presumed land plants add support to this model. In addition, such spore types suggest that the temporal relationship of meiosis to sporopollenin deposition was less strictly controlled than in extant land plants.

The striped colour pattern and striped/non-striped polymorphism in snakes (reptilia: ophidia)*

Abstract: Summary The occurrence of striped colour patterns and of striped/non-striped polymorphism systems among snakes is reviewed from literature data augmented by some personal observations. Among 1367 species, 190 were striped or had striped morphs. Of 11 families, the striped pattern was common mainly among Colubridae, presumably in relation to the active escape behaviour strategy, prevalent in this family. The striped species tended to cluster in a small number of genera. The 40 striped/non-striped polymorphism systems found, fall into five categories, according to the coloration patterns of the alternative morphs: (I) blotched (cryptic); (2) barred (or ringed); (3) plain; (4) melanistic; (5) albinistic. Most polymorphisms are presumably maintained by eco-behavioural trade-offs, depending on the category and on the habitat: The striped morph is presumed more effective in active escape and sometimes also in camouflage; the alternative morph may be more effective in camouflage, in active escape or in thermoregulation. Hence morph frequency depends on the habitat. Striped-albinistic polymorphism in Elaphe climacophora presumably depends on human protection of the albino morph.

Alternation of generations in algae: its complexity, maintenance and evolution

Abstract: Summary Life histories of photosynthetic eukaryotes traditionally-termed algae exhibit a considerably greater degree of complexity than those of ‘higher cryptogams.’ Some algae have a so-called ‘obligate’alternation between spore-producing and gamete-producing phases, but the majority seem capable of following other pathways depending upon environmental conditions. In only four algal classes do life histories show a change in morphological and/or nuclear phases. The following basic life histories are recognized in the Chlorophyceae, Phaeophyceae and Rhodophyceae:(a) monophasic, a diploid or haploid phase, (b) two or more phases, most commonly an alternation of an isomorphic or heteromorphic haploid gametangial phase and a diploid sporangial phase, and (c) three phases (unique to florideophyte Rhodophyceae), with a diploid spore-producing phase (carposporophyte) developing on the gametangial phase, a diploid phase (tetrasporophyte if meiosis is sporic) and a haploid gametangial phase. Evidence from recent research indicates that in many algae there is an uncoupling of the morphological and nuclear phases. The dominance of one phase and suppression of another has been suggested to be due to the common occurrence in algae of apogamy, apomeiosis and parthenogenesis. Free-living morphs in heteromorphic life histories may be morphologically so dissimilar that formerly they were attributed to different genera. Evolution of the carposporangial phase in red algae is speculated to be a means of achieving zygotic amplification to compensate for the infrequency of syngamy. Such amplification allows the production of a large number of dispersible products from a single fertilization. The direct development of a free-living tetrasporangial phase is considered another mechanism for achieving amplification. In freshwater red algae the growth of an upright phase from a perennial microscopic one is considered an adaptation for maintaining their upstream position. Life history pathways in algae are controlled by subtle environmental influences (e.g. photoperiodism, temperature, light quality, nutrients). Experimental evidence is lacking to support the contention that spatial and/or temporal partitioning of the environment is a mechanism favouring the maintenance of heteromorphy. Herbivory is known to be an important selective force suppressing some morphs and accentuating the seasonal dominance of others. Differential resistance of morphs to herbivory in environments where grazing intensity is predictable may lead to the selective maintenance of heteromorphy. Algal life history patterns are unexplored in terms of evolutionary processes. Various models for the evolution of biphasic or polyphasic life histories stress the importance of the capacity for both asexual dispersal of successful genotypes and for the generation of new genotypes via meiosis and syngamy. All evidence points to the fact that many life history processes operative in algae differ significantly from those described for other cryptogams.

Rhythm, periodicity and polarity as bases for morphogenesis in plants

Abstract: CONTENTS

The regulation of alternation of generation in flowering plants

Abstract: Summary The developmental changes involved in the alternation of generation represent the major gene-switching events in the life history of plants. While a large number of genes are common to both sporophyte and gametophyte, many thousand sequences are specifically expressed in each generation; indeed, certain key constituents (e.g. tubulin) are encoded by different genes in each generation, indicating that sporophyte and gametophyte are responding to different evolutionary pressures. Evidence is accumulating that major gene-switching events in plants, such as flowering, are regulated by complex control systems which ensures that development occurs only in the correct groups of cells at the appropriate time. A similar, or more sophisticated system might thus be expected to regulate alternation of generation. It is not possible to manipulate alternation of generation in a similar fashion to flowering, but study of apparent aberrations of development occurring in nature and in vitro suggests that alternation only occurs in cells which have become competent to receive particular developmental stimuli. Further, in certain cases, competent cells may be switched either into sporophytic or gametophytic developmental pathways depending upon the nature of the stimulus. Acquisition of competence seems to involve isolation of cells from the symplast, some cytoplasmic dedifferentiation, and perhaps cell cycle arrest or transition. The stimuli in vivo appear metabolic in nature, although embryogenesis may be activated by specific classes of glycoproteins. Interestingly, examination of agamospermic systems suggests that fertilization of the egg per se is not the signal which activates sporophytic development. Once competent cells have received the stimulus they start to develop, with no delay in a ‘determined’ state. Sporophytic and gametophytic development in vivo and in vitro both start with an asymmetric division, except for the female gametophyte which may arise via a range of developmental pathways, depending on the species.

Alternation of generations in ferns: mechanisms and significance

Abstract: Summary Arguments against the compiling of generalized life cycles summarizing alternation of generations in ferns are presented, and some common misconceptions about breeding systems addressed. What little is known or can be deduced about time frames, mechanisms and significance of alternation events in the lives of two species: bracken fern (Pteridium) and Killarney fern (Trichomanes speciosum) is presented. Evidence is provided that gametophytes may play a more important role in survival of both these species than previously suspected, and the need for more long-term studies and experiments/measurements of ferns in natural conditions/populations is stressed.

Synthesis of a self‐contained concept of the molecular mechanism of energy interconversion by h+‐transporting atp synthase

Abstract: The original aim of the review has been to probe into the validity of the paradigm on the high energy-carrier function of ATP. It seemed to be called into question on the basis of findings with H(+)-transporting ATP synthase suggesting the formation of ATP from ADP and Pi without energy input. Thus, ATP appeared as a low-energy compound. Starting from the current, rich knowledge of the molecular structure and the inviting thinking on the mechanism of H(+)-transporting ATP synthase, we have endeavoured to freshly interpret and integrate the pertinent observations in the light of the comprehensively derived model of the molecular mechanism of energy interconversion by Na+/K(+)-transporting ATPase. In this way, we have uncovered the common mechanistic elements of the two energy-interconverting enzymes. The emerging purpose of the present paper has been the 'synthesis' of a self-contained concept of the molecular mechanism of the interconversion of electrochemical and chemical Gibbs energies by H(+)-transporting ATP synthase. The outcome is reflected in the following tentative evaluations. 1. In ATP hydrolysis, the great Gibbs energy change which is observed in solution, is largely conserved by the F1 sector of ATP synthase as mechanical Gibbs energy in the enzyme's protein fabric, so that it can be utilized in the resynthesis of ATP from enzyme-bound ADP and Pi. The plainly measured low Gibbs energy change results from large compensating enthalpy and entropy changes that reflect the underlying changes in protein conformation. 2. In stoichiometric ATP synthesis by F1 sector from ADP and Pi bound to the catalytic centre, their intrinsic binding energy brings about a loss of peptide chain entropy that makes possible an entropy-driven ATP formation. 3. The driving force for ATP synthesis cannot be the high Gibbs energy change on binding of product ATP; the tight ATP-enzyme complex rather is a low Gibbs energy intermediate from which escape is difficult. 4. The catalytic centre exists either in an open state unable to firmly bind the substrate-product couple, or in a closed state protecting formed ATP from facile hydrolysis by ambient water. 5. The cleft closure, induced by binding of Pi and ADP or ATP, does not necessarily need external energy supply, because the cleft closure proceeds from rigid domain rotations which can occur rather spontaneously. In further analogy to adenylate kinase, the driving force of this domain movement presumably comes from the electrostatic interactions between phosphate moieties and arginine side chains in the catalytic centre.(ABSTRACT TRUNCATED AT 400 WORDS)