Showing papers in "Biological Reviews in 1990"

Fluctuating asymmetry: an epigenetic measure of stress

Abstract: (1) Fluctuating asymmetry (FA) is a useful trait for monitoring stress in the laboratory and in natural environments. (2) Both genomic and environmental changes can increase FA which represents a deterioration in developmental homeostasis apparent in adult morphology. Genetic perturbations include intense directional selection and certain specific genes. Environmental perturbations include temperature extremes in particular, protein deprivation, audiogenic stress, and exposure to pollutants. (3) There is a negative association between FA and heterozygosity in a range of taxa especially fish, a result consistent with FA being a measure of fitness. (4) Scattered reports on non-experimental populations are consistent with experiments under controlled laboratory conditions. FA tends to increase as habitats become ecologically marginal; this includes exposure to environmental toxicants. (5) In our own species, FA of an increasing range of traits has been related to both environmental and genomic stress. (6) Domestication increases FA of the strength of homologous long bones of vertebrate species due to a relaxation of natural selection. (7) FA levels are paralleled by the incidence of skeletal abnormalities in stressful environments. (8) Increased FA is a reflection of poorer developmental homeostasis at the molecular, chromosomal and epigenetic levels.

Hatching asynchrony in altricial birds

Abstract: Summary 1. The review aims to provide a simple conceptual framework on which to place recent studies of hatching asynchrony in altricial birds and to assess the evidence used in support of specific hypotheses. 2. Hatching asynchrony arises bsecause parents start incubation before laying is complete, but the precision of parental control is largely unknown. 3. Hypothesses concerning the functional significance of hatching asynchrony fall into four broad types. Hatching asynchrony might: (i) arise because of selection on the timing of events during the nesting period; (ii) facilitate the adaptive reduction in brood size; (iii) increase the energetic efficiency of raising the brood, or (iv) result from environmental or phylogenetic constraints. 4. The incubation pattern could function to minimize the losses of eggs, nestlings or adults to predators (or climatic sources of mortality), particularly in species which cannot actively defend their nest. The best evidence comes from comparative studies of hatching asynchrony. Early incubation might also be favoured if the food supply declines sharply through the breeding season, although the evidence is weak and indirect, or if there is a risk of brood parasitism. In species in which only the female incubates, early incubation could ‘force’ the male to invest more in the nestlings, but this idea remains to be tested. Males may be constrained by the risk of cuckoldry to delay incubation until laying is complete. 5. Hatching asynchrony could be adaptive by enabling the efficient reduction of brood size if food proves short after hatching (primarily because of a shortage of food in the environment or possibly because of a large proportion of ‘expensive’ nestlings in the brood in species which are sexually dimorphic). Observational evidence is often consistent with this hypothesis but few experimental studies provide adequate tests. Brood reduction could be adaptive in species (primarily eagles and pelecaniformes) which lay an extra egg to act as insurance against hatching failure, and again hatching asynchrony might facilitate brood reduction, although there are few experimental tests on such species. Hatching asynchrony might also enable sex ratio manipulation through selective brood reduction, although there is as yet no clear supportive evidence. 6. Ins species in which young have a marked peak in energy demand during the period of parental care, hatching asynchrony can reduce the peak demand of the brood, which might allow the parents to raise more healthy young. In many species such savings are likely to be small or absent. There is some behavioural evidence that hatching asynchrony can reduce fighting amongst nestlings and therefore lead to the more efficient use of energy by the brood. In general this effect seems small and the only energetic study found no difference in the energy requirements of synchronous and asynchronous broods. Other possible energetic advantages to hatching asynchrony have not been tested. 7. Environmental conditions during laying can influence both egg size and laying interval in aerial insectivores, and might directly influence incubation in this and other groups. Thus some variation in hatching asynchrony and the relative size of siblings is probably non-adaptive. The variability of incubation pattern within and across species suggests that hatching asynchrony is not under strong phylogenetic constraint. 8. The hypotheses about the adaptive significance of hatching asynchrony are complementary rather than mutually exclusive: within a species, several selective pressures could influence the optimal incubation pattern, and the relative importance of selective pressures will differ among species. Furthermore one should expect that the incubation pattern and parent–offspring interactions will be coadapted to maximize brood productivity.

Development and evolutionary origins of vertebrate skeletogenic and odontogenic tissues.

Abstract: This review deals with the following seven aspects of vertebrate skeletogenic and odontogenic tissues. 1. The evolutionary sequence in which the tissues appeared amongst the lower craniate taxa. 2. The topographic association between skeletal (cartilage, bone) and dental (dentine, cement, enamel) tissues in the oldest vertebrates of each major taxon. 3. The separate developmental origin of the exo- and endoskeletons. 4. The neural-crest origin of cranial skeletogenic and odontogenic tissues in extant vertebrates. 5. The neural-crest origin of trunk dermal skeletogenic and odontogenic tissues in extant vertebrates. 6. The developmental processes that control differentiation of skeletogenic and odontogenic tissues in extant vertebrates. 7. Maintenance of developmental interactions regulating skeletogenic/odontogenic differentiation across vertebrate taxa. We derive twelve postulates, eight relating to the earliest vertebrate skeletogenic and odontogenic tissues and four relating to the development of these tissues in extant vertebrates and extrapolate the developmental data back to the evolutionary origin of vertebrate skeletogenic and odontogenic tissues. The conclusions that we draw from this analysis are as follows. 8. The dermal exoskeleton of thelodonts, heterostracans and osteostracans consisted of dentine, attachment tissue (cement or bone), and bone. 9. Cartilage (unmineralized) can be inferred to have been present in heterostracans and osteostracans, and globular mineralized cartilage was present in Eriptychius, an early Middle Ordovician vertebrate unassigned to any established group, but assumed to be a stem agnathan. 10. Enamel and possibly also enameloid was present in some early agnathans of uncertain affinities. The majority of dentine tubercles were bare. 11. The contemporaneous appearance of cellular and acellular bone in heterostracans and osteostracans during the Ordovician provides no clue as to whether one is more primitive than the other. 12. We interpret aspidin as being developmentally related to the odontogenic attachment tissues, either closer to dentine or a form of cement, rather than as derived from bone. 13. Dentine is present in the stratigraphically oldest (Cambrian) assumed vertebrate fossils, at present some only included as Problematica, and is cladistically primitive, relative to bone. 14. The first vertebrate exoskeletal skeletogenic ability was expressed as denticles of dentine. 15. Dentine, the bone of attachment associated with dentine, the basal bone to which dermal denticles are fused and cartilage of the Ordovician agnathan dermal exoskeleton were all derived from the neural crest and not from mesoderm. Therefore the earliest vertebrate skeletogenic/odontogenic tissues were of neural-crest origin.(ABSTRACT TRUNCATED AT 400 WORDS)

Patterns in the geographical ranges of species

Abstract: Summary 1. A growing number of studies have documented patterns in species geographic ranges relevant to the study of community structure. These include patterns in the frequency of geographic ranges of different sizes, and in the interaction of range sizes with population abundances and variabilities, body sizes, trophic characteristics and extinction probabilities. 2. Agreement between hypothesized and observed relationships is reasonably good, but we do not know how general the patterns are. Analysis of patterns has focused upon a few taxonomic groups, a bias largely resulting from a lack of information on the geographic distributions of most taxa. 3. Many of the patterns are interrelated, and although theoretical bases to all the patterns can be suggested, it is possible that some are artifacts. 4. Taylor power plots give us a means of making predictions about population behaviour as it pertains to geographic ranges. Some of these predictions suggest that previous conceptions of such interactions have been too narrow, but empirical analyses of these patterns will be hampered by the difficulty of measuring population variability. 5. In general, our knowledge of the structure, and spatial and temporal behaviour, of species geographic ranges remains poor.

Sex ratios and conditions required for environmental sex determination in animals

Abstract: Conclusions and Summary Environmental sex determination (ESD) is a system of sexual determination that is influenced by a variable environment. Once sex is determined it is then fixed for life. The model of Charnov & Bull (1977) proposes that ESD is favoured by natural selection when an individual's fitness as a male or female is strongly influenced by environmental conditions and when the individual has little control over which environment it will experience. Adaptive sex ratio variation is considerably easier for organisms with ESD, and this feature is the ultimate cause for the evolution and maintenance of ESD. ESD is taxonomically widely expressed, and more cases are likely to be discovered. Both environmental and genotypic sex determination mechanisms are found in closely related species. Evidence of geographical variation in the degree and in the critical environmental values of ESD within the same species has also been discovered, e.g. in the fish Menidia menidia and in the crustacean Gammarus duebeni. The factors causing sex determination in invertebrates include temperature, daylength, nutrition, density, humidity, ionic composition of the environment, pH, carbon dioxide, UV light, metabolic products, parasites, exposure to the opposite sex of the same species, and in parasitoids also host size, age and type. In vertebrates temperature is the dominant factor causing sex determination, though in fish also pH, salinity, light, water quality and nutrition, and in turtles water potential of the substrate have some effect on the sex expression. Most of these factors influence growth through resource availability or developmental speed. In most cases of ESD in invertebrates and fish, the environmental factor has a gradual effect on the sex expression, in contrast to the typical steep threshold mode found in reptiles. These differences might be due to the fact that invertebrates exhibiting ESD are commonly parasitic or confined to aquatic environments, where less spatial microhabitat differentiation exists. Sex ratio data available from nature for animals with ESD are quite limited, except for reptiles. In the laboratory sex ratios can be varied more widely than what is observed in nature. There are a number of characteristic features some of which are found in each species exhibiting ESD: (1) Patchy environments, (2) variable sex ratios, (3) parthenogenesis in addition to bisexuality, (4) parasitism, (5) aquatic habitats, (6) sexual dimorphism, (7) females larger than males, and (8) local mate competition.

Mechanisms for the control of gene activity during development

Abstract: (4) Somatic cells of Drosophila (5) Imprinting and germ line transmission (6) Maize transposable elements . . . . . . . . . . . (7) Modification of integrated DNA in fungi (8) Cancer and ageing . . . . . . . . . . . . . V . Conclusions . . . . . . . . . . . . . . . ( I ) General significance of DNA methylation (2) Major components of development . . . . . . . . . . VI . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . .

Ovariole types and the phylogeny of hexapods

Abstract: CONTENTS

Chemical communication in insect communities: a guide to insect pheromones with special emphasis on social insects

Abstract: SUMMARY 1Chemical communication plays an important part in the lives of insects, and particularly in lives of those that live in groups or social organizations. 2Chemicals which are used in communication in the general sense are called semiochemicals, and there are a number of subdivisions recognized under this title. 3Pheromones are a category of semiochemicals which are used for communication between individuals of the same species. 4Pheromones are in turn subdivided into primer and releaser pheromones. The former produce a relatively long-lasting physiological change in the receiver, and the latter stimulate the receiver to some immediate behavioural response. 5Far more is known about releaser pheromones at present because they are easier to study. 6Nine categories of releaser pheromone are recognized here, used by both social and non-social insects. 7Sex pheromones are widely used to bring the sexes together for mating, and they have been extensively studied in Lepidoptera. 8Invitation pheromones, encouraging the species to feed or oviposit at an explored site, are not extensively known. 9Aggregation pheromones are designed to bring individuals together into groups which may be temporary in sub-social insects, or permanent in social insects. 10Dispersal or spacing pheromones are used by other species to reduce intraspecific competition for scarce resources. 11Alarm pheromones are a broad and sometimes unclearly defined group which communicate alarm or attack, chiefly in colonial species. 12Trail pheromones, applied to a surface by an individual, to be followed by another, are confined to Hymenoptera, Isoptera and a few Lepidoptera as far as is known. 13Territorial and home range pheromones may be widely distributed, but as yet few of them have been recognized. 14Surface and funeral pheromones are even less well known. Surface pheromones may play a large part in species or colony recognition. 15We can expect the number and complexity of pheromones to be much greater in social insects, a part of the subject which until now has received relatively less attention. 16As our understanding of the subject grows we may expect other categories to be added to this list.

Early trends in the evolution of tribosphenic molars

Abstract: . . . . . . . . . . . . . . . I . Introduction 529 11. Nomenclature and mensuration 531 111. Peramuridae 534 IV. Early Cretaceous tribosphenids 535 V. Deltatheridiids and possible relatives 538 VI. Marsupials 5 40 VII. Placentals 541 VIII. The placental radiation 544 IX. The Early Cretaceous radiation X. Acknowledgements 549 XI. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 547

The evolution of reversed sexual dimorphism in size in monogamous species of birds

Abstract: Summary Reversed sexual dimorphism in size (RSD) occurs in most species of several taxonomic groups of birds. The hypotheses proposed to explain this phenomenon are examined theoretically, using inequalities to state selection in the most rigorous possible terms. The most pertinent empirical evidence is also examined critically. Proponents of hypotheses on the evolution of RSD have failed to consider the genetic constraints on the evolution of dimorphism. Selection for dimorphism can act on only that small portion of the genetic determination of body size that is sex limited. In general, selection for body size is much more likely to lead to a similar change (e.g. larger) in both sexes than to dimorphism. The most popular hypotheses involve selection for size-related differences in foraging ability. It is unlikely that there is variation in size-related foraging differences available for selection in a monomorphic, ancestral population. Foraging differences between the sexes cannot lead to the evolution of RSD; evolution of large and small morphs of both sexes is a more likely outcome. Selection for sex-role differentiation factors (e.g. large females lay larger eggs, small males are more agile in flight) can lead to the evolution of RSD, but only if the magnitudes of opposing selection for small males and for large females are equal. Combining selection for size-related foraging differences with selection for sex-role differentiation factors hinders the evolution of RSD until the sexes differ in size by 3 s.d. Empirical evidence supports this assertion: statistically significant differences between the sexes in the size of prey taken are found only in highly dimorphic species. The sex-role differentiation factors that have been proposed appear unlikely to provide the equal selection necessary for the evolution of RSD. Several authors have proposed that small size in males is selected for foraging ability and large size in females for some sex-role differentiation factor. Males cannot be more efficient foragers without females being less efficient and efficiency cannot be a factor only when the male is feeding his family. RSD cannot evolve in monogamous species if large females survive less well than small males. RSD might evolve as the result of sexual selection for small size in males and constraints on the reduction of size in females because of some factor associated with reproduction. Examination of seven studies indicating a relationship between female size and reproductive success shows very little unequivocal evidence for small size in females allowing breeding earlier in the season. Large size in females allows females to breed at a younger age in the sparrowhawk and pairs to form more rapidly in three species of sandpipers. Both of these may be the result of sexual selection. There are fewer theoretical problems with sexual selection as a cause for the evolution of RSD than with the other hypotheses. Empirical evidence for sexual selection is scarce but better than that for the other hypotheses. Evidence is contradictory for the selection of small size in males for agility in aerial displays for courtship or defence of territory. Large size in females does not appear to be the result of selection for competitive ability to obtain mates. Facilitation of female dominance and hence of the formation and maintenance of a pair bond is the most viable explanation of the evolution of RSD. It is most likely that all dimorphism (normal or reversed) is the result of sexual selection. RSD is correlated with birds in the diet in the Falconiformes and this is a central theme in the foraging hypotheses. This correlation may be because birds are abundant and available in a continuum of sizes, thus permitting but not causing the evolution of RSD or because species that prey upon birds are better equipped physically (and perhaps more likely behaviourally) to inflict damaging attacks on conspecifics and the greater RSD increases female dominance and the ease of pair formation.

Quantitative variations of nuclear dna during plant development: a critical analysis

Abstract: SUMMARY In this review a critical analysis is made of the quantitative DNA variations that take place during plant development. These include underreplication, loss and amplification of repeated sequences; particular attention is devoted to quantitative variations of non-coding DNA sequences. Examples reported in the literature are analysed in relation to the different biological phenomena they accompany such as vegetative development, phase changes, senescence, dedifferentiation or tumour formation. The author also considers the influence that external elements such as growth factors, chemical substances, culture medium composition, or, also, different stimuli of environmental character, can have on these quantitative variations of nuclear DNA sequences. Finally the different hypotheses on the role that repetitive DNA sequences (particularly those that are highly repeated) can have on the life of the cell are analysed and discussed. The experimental evidence, although fragmentary and sometimes contradictory, leads to the conclusion that the plant genome is not so ‘quantitatively static’ as was once hought, but is instead characterized by a dynamic plasticity; the role of repeated DNA sequences in the plant development seems to be much more active, than the role hypothesized by many authors in the past.

The evolution of heteromorphic sex chromosomes.

Abstract: SUMMARY The facts and ideas which have been discussed lead to the following synthesis and model. 1 Heteromorphic sex chromosomes evolved from a pair of homomorphic chromosomes which had an allelic difference at the sex-determining locus. 2 The first step in the evolution of sex-chromosome heteromorphism involved either a conformational or a structural difference between the homologues. A structural difference could have arisen through a rearrangement such as an inversion or a translocation. A conformational difference could have occurred if the sex-determining locus was located in a chromosomal domain which behaved as a single control unit and involved a substantial segment of the chromosome. It is assumed that any conformational difference present in somatic cells would have been maintained in meiotic prophase. 3 Lack of conformational or structural homology between the sex chromosomes led to meiotic pairing failure. Since pairing failure reduced fertility, mechanisms preventing it had a selective advantage. Meiotic inactivation (heterochromatinization) of the differential region of the X chromosome in species with heterogametic males and euchromatinization of the W in species with heterogametic females are such mechanisms, and through them the pairing problems are avoided. 4 Structural and conformational differences between the sex chromosomes in the heterogametic sex reduced recombination. In heterogametic males recombination was reduced still further by the heterochromatinization of the X chromosome, which evolved in response to selection against meiotic pairing failure. 5 Suppression of recombination resulted in an increase in the mutation rate and an increased rate of fixation of deleterious mutations in the recombination-free chromosome regions. Functional degeneration of the genetically isolated regions of the Y and W was the result. In XY males this often led to further meiotic inactivation of the differential region of the X chromosome, and in this way an evolutionary positive-feedback loop may have been established. 6 Structural degeneration (loss of material) followed functional degeneration of Y or W chromosomes either because the functionally degenerate genes had deleterious effects which made their loss a selective advantage, or because shorter chromosomes were selectively neutral and became fixed by chance. 7 The evolutionary routes to sex-chromosome heteromorphism in groups with female heterogamety are more limited than in those with male heterogamety. Oocytes are usually large and long-lived, and are likely to need the products of X- or Z-linked genes. Meiotic inactivation of these chromosomes is therefore unlikely. In the oocytes of ZW females, meiotic pairing failure is avoided through euchromatinization of the W rather than heterochromatinization of the Z chromosome. Since both chromosomes are euchromatic, recombination should occur. There is therefore no reason for the functional or structural degeneration of the W unless it is initiated by a reduction in crossing-over brought about by a structural change such as a paracentric inversion. A conformational difference between the homologues could have led to functional degeneration of the W only if meiotic pairing and recombination were uncoupled as they are in the Lepidoptera. 8 Sex-chromosome heteromorphism usually causes gene-dosage differences between males and females. In some organisms with male heterogamety, the mechanisms controlling meiotic X-inactivation in the spermatocyte were modified in ways which led to dosage compensation in the somatic cells. Since no meiotic inactivation mechanisms occur for Z chromosomes, dosage-compensation systems comparable to those found in species with male heterogamety could not evolve in species with femal heterogamety. 9 In marsupials, dosage compensation is brought about by preferential inactivation of the paternal X chromosomes in females. The general suppression of recombination observed in female marsupials could be a consequence of selection pressures resulting from preferential X-inactivation. 10 Sex-chromosome heteromorphism and the conformational changes of meiosis which are associated with it may be the reason why X chromosomes show more genomic imprinting than autosomes. The asymmetrical pattern of transmission of sex chromosomes may give parental imprinting a selective advantage. 11 Genetic changes affecting the conformation of sex chromosomes may be important causes of the sterility and inviability found in hybrids between individuals from populations which have diverged during periods of isolation.

Autoxidation of oxymyoglobin: a meeting point of the stabilization and the activation of molecular oxygen.

Abstract: Summary 1 The primary events of haemoprotein reactions with molecular oxygen have been re-examined by placing special emphasis upon the reduction properties of dioxygen. 2 In the stepwise reduction of O2 to water via hydrogen peroxide, the addition of the first electron is an unfavourable, uphill process with the midpoint potential of –0.33 V, all the subsequent steps being downhill. This thermodynamic barrier to the first step is, therefore, a most crucial ridge located between the stabilization and the activation of dioxygen performed by haemoproteins. 3 If the proteins have a redox potential much higher than –0.33 V, molecular oxygen must bind to the proteins stably and reversibly. In Mb or Hb, however, the FeO2 centre is always subject to a nucleophilic attack of the water molecule or hydroxyl ion, which can enter the haem pocket from the surrounding solvent. These can cause irreversible oxidation of the FeO2 bonding to the ferric met-form with generation of the superoxide anion. 4 In cases of the oxygen activation, if haemoproteins have a redox potential lower than or close to –0.33 V, the first reduction of O2 to O−2 would be a spontaneous process. Cytochrome P-450 provides such an example and can facilitate the subsequent addition of electrons that leads to the breaking of the O–O bond to yield the hydroxylating species. 5 As to the proteins whose redox potential is not facilitative and appreciably higher than –0.33 V, a bimetallic, concerted, two-equivalent reduction of the bound dioxygen to the peroxide level would be much more favoured without the intermediate formation of O−2. This is probably the case of cytochrome c oxidase for the reduction of O2 to water. 6 The redox potential diagrams thus visualize various aspects of the ways haemoproteins overcome their thermodynamic constraints and carry out their specific functions in the stabilization and the activation of molecular oxygen.

New targets for cancer chemotherapy – poly(adpribosylation) processing and polyisoprene metabolism*

Abstract: Advances in the study of cellular metabolism are beginning to divert necessary attention away from DNA as the conventional (and direct) target for cancer chemotherapeutic agents. Accordingly, this paper presents alternative targets, and puts forward suggestions for certain appropriately designed drugs whose effects may provide a basis for a more pharmacological approach to cancer chemotherapy. This involves consideration of the action of anti-tumour N6-substituted adenine ribonucleosides, principally by reason of their evident effects both on the poly(ADP-ribosylation) processing of nuclear proteins and through polyisoprene metabolism.

Developmental heterochrony in ciliated protozoa: overlap of asexual and sexual cycles during conjugation

Abstract: CONTENTS

Plant growth and development in molecular perspective

Abstract: Summary 1 After some false starts in which inactive plant substances were isolated, the isolation and identification of auxin as the growth substance at the meristems and of ethylene as the ripening agent in climacteric fruits represented outstanding achievements. 2 In early work, the non-localized origin of auxin at the meristem and its possible transport for coleoptile development were obscured by the superimposition on the results of physiological experiments of the idea of a close parallelism between the plant-growth substances and mammalian hormones. At that time, an absence of chemical instrumentation, suitable for measurement of the tissue levels, compounded the difficulty in interpreting available physiological evidence. 3 Member(s) of each of the five groups of naturally occurring plant-growth substances, namely the auxins, cytokinins, gibberellins, ethylene and the growth inhibitors, including abscisic acid, are biologically active at a concentration of 10 μm or less, however, and in this respect they would appear to qualify as candidate phytohormones. 4 The sensitivity of plant cells to phytohormones contributes to plant growth and development, and both the variations in sensitivity, for example, of wheat coleoptiles towards growth and the growth of the coleoptiles per se give parallel unimodal relationships with regard to time; the curve representing sensitivity precedes that for growth. A new graphical analysis implies that the growth sensitivity and growth rate functions are mutually interdependent. 5 The assumption is made in point 4 that growth substance complexes with receptor protein in growth-sensitive cells, and the concept of receptors would provide explanation for the obvious amplification of effects induced by growth substances. 6 Numerous biological situations occur in which the presence of significant amounts of plant hormone controls growth and development. In gravitropism and phototropism, tropistic curvature depends on the difference in physiological concentration of auxin on the two sides of the organ concerned. In infected tobacco plants, the cytokinin to auxin ratios for the tumours determine the kind of development (tumours and shoots, tumours only or tumours plus roots), which takes place. 7 Auxin-binding protein has been identified immunologically, and isolated. Work with hormone receptors for gibberellin does not afford unequivocal evidence for more than one primary site of action. Hitherto, no specific receptor protein is known for cytokinins. 8 Clear evidence derives, both from structure–activity relationships and from unimodal concentration–response curves, for receptor specificity to auxin action. There is also evidence for a structure–activity relationship in respect of the cytokinin series of compounds. 9 From the evidence (points 1–8), there emerges a picture of hormone-induced growth and development of plant cells, which have been made sensitive to hormone through the presence of specific receptor proteins. 10 That plant growth and developmental processes involve changes in gene expression would seem to follow from the totipotent nature of meristematic cells, which are capable of specialization in response to phytohormones. 11 Auxin regulates de novo synthesis of mRNAs encoding polypeptides essential to the auxin-induced early process of cell elongation. In fact, auxin regulates the concentrations of several authenticated mRNAs and proteins, for example, in elongating soyabean hypocotyl sections. 12 Furthermore, two cDNA clones, termed pJCW1 and pJCW2 have been isolated with the properties expected of mRNAs involved in the rate-limiting stage of cell elongation. The evidence suggests that the change in relative abundance of the JCW1 and JCW2 RNAs is an obligatory auxin-dependent response. Hence, the action of cytokinin in auxin-induced cell elongation would seem to be concerned with the inhibition of rate-limiting proteins, and in fact cytokinin inhibits protein synthesis in excised soyabean hypocotyl. 13 Biosystemic experiments on some rapid effects of synthetic auxin growth regulators on mRNA levels in vitro show that there is only partial similarity between those found in pea and soyabean spp. (Leguminosae). 14 Two identified sequences, namely TGATAAAAG and GGCAGCATGCA, of two auxin-regulated soyabean genes afford a means for determining whether the auxin-regulation of expression of these genes involves trans-acting regulatory factors. 15 The obligatory auxin-induced responses with regard to cell elongation and growth (q.v.) would seem to precede the somewhat mechanical growth properties by which auxin receptive cells secrete H+ and lower the pH to yield increased cell-wall plasticity. 16 In vertically oriented soyabean seedlings, auxin-regulated RNAs are distributed symmetrically in the elongating region of the hypocotyl, whereas in horizontally oriented seedlings the distribution becomes asymmetrical within a few minutes of horizontal gravitational stimulation. The dynamic expression of auxin-regulated genes is related to the morphogenetic response, initiated by re-distribution of endogenous auxin (point 6). 17 In the germination of seeds, the mobilization of food reserves requires hydrolytic enzymes and, in barley grains, gibberellic acid induces de novo biosynthesis of α-amylase and protease. The genetic implications are discussed, and the requirement of dicotyledonous and gymnospermous seeds for the presence of gibberellins is explored. 18 In ripening climacteric fruits, ethylene-induced change(s) in gene expression cause de novo biosynthesis of polygalacturonase, which degrades the cell-wall pectin fraction. 19 Accordingly, incontestible evidence has been mustered for the proposition that hormone-regulated plant growth and development involves hormone-regulated gene expression. 20 As well as the phytohormones, certain environmental factors, such as white light and stress (including anaerobiosis, chilling, heat shock, heavy metal exposure, u.v. light and wounding) have the capacity to regulate gene expression in plants at important stages in growth and development. Discussion at the genetic level focuses on changes produced by: 21 The synthesis of phytohormones is significant. For example, as u.v. light-induced regulation of genes produces enzymes for auxin synthesis, it may be responsible in seeds for the endospermal generation of auxins, concerned with the epicotyl/hypocotyl growth in the seedlings. 22 Hormones and certain environemental factors (q.v.) initiate some of the numerous stages in plant growth and development, but the regulatory factors are obscure in some other biological situations, such as: 23 Utilization of an appropriately re-constituted plant DNA polymerase i in vitro system might enable the type and frequency of misincorporation, produced by plant-growth factors, to be studied. Base-pair substitution changes were produced in strains of crop plant, made resistant to a specific herbicide by genetic engineering (see Hathway, 1989). It is feasible that auxin may behave as a reagent in the chemical sense to effect intramolecular change(s) in some of the sequences concerned, leading to the frame-shift changes observed (see Ainley et al., 1988).

Embryonic resistance to chemical and physical factors: manifestation, mechanism, role in reproduction and in adaptation to ecology

Abstract: Chemical and physical factors may adversely affect embryonic development. As an example of chemical factors, the effects of diabetic metabolic factors on embryonic development in mammals was reviewed. The existence of a stage-dependent reaction of embryos was found. At preimplantation stages diabetic metabolic factors are embryotoxic and lethal, and the blastocysts reacted by an "all-or-none" response. Early somite embryos showed a higher resistance to the effects of diabetic metabolic factors resulting in various types of malformations. Both groups of embryos showed a very high sensitivity to the effects of combined diabetic metabolic factors. Congenital defects in term foetuses were lower than those observed during middle phases of pregnancy because some of the severely malformed embryos resorb during gestation. The effects of temperature on embryonic development were presented as an example of physical influences. In man, hyperthermia in pregnancy seems to correlate with defects in the development of the nervous and skeletal systems. In domestic animals, changes in environmental temperature correlated with depressions of reproduction rate. In laboratory animals, hyperthermia caused the development of congenital malformations. Stage-dependent as well as genetic differences in embryonic susceptibility to hyperthermia were found. Critical periods in sensitivity of embryos to hyperthermic influences were also observed. It has been shown that, in spite of similar external manifestations of the reaction of embryos to effects of diabetes and hyperthermia, the mechanism of these reactions was different. High resistance of early reptile and bird embryos to influences of temperature was considered as an example of morphofunctional adaptations in early embryogenesis of vertebrates to their development in terrestrial conditions.