Plasticity in cultured carotid body chemoreceptors: Environmental modulation of GAP-43 and neurofilament

In this study we use dissociated cell cultures of the rat carotid body to investigate the adaptive capabilities of endogenous oxygen chemoreceptors, following chronic stimulation by various environmental factors. These oxygen chemoreceptors are catecholamine-containing glomus cells, which derive from the neural crest and resemble adrenal medullary chromaffin cells. Using double-label immunofluorescence, we found that chronic exposure of carotid body cultures to hypoxia (2% to 10% oxygen) caused a significant fraction of tyrosine hydroxylase-positive (TH+) glomus cells to acquire detectable immunoreactivity for growth-associated protein gap-43. The effect was dose-dependent and peaked around an oxygen tension of 6%, where approximately 30% of glomus cells were GAP-43 positive. Treatment with agents that elevate intracellular cyclic adenosine monophosphate (cAMP) (i.e., dibutyryl cAMP or forskolin) also markedly stimulated GAP-43 expression. Since hypoxia is known to increase cAMP levels in glomus cells, it is possible that the effect of hypoxia on GAP-43 expression was mediated, at least in part, by a cAMP-dependent pathway. Unlike hypoxia, however, cAMP analogs also stimulated neurofilament (NF 68 or NF 160 kD) expression and neurite outgrowth in glomus cells, and these properties were enhanced by retinoic acid. Nerve growth factor, which promotes neuronal differentiation in related crest-derived endocrine cells, and dibutyryl cGMP were ineffective. Thus, it appears that postnatal glomus cells are plastic and can express neuronal traits in vitro. However, since hypoxia stimulated GAP-43 expression, without promoting neurite outgrowth, it appears that the two processes can be uncoupled. We suggest that stimulation of GAP-43 by hypoxia may be important for other physiological processes, e.g., enhancing neurotransmitter release or sensitization of G-protein–coupled receptor transduction. © 1995 John Wiley & Sons, Inc.     

Matching field and laboratory environments: effects of neglecting daily temperature variation on insect reaction norms

The tropical butterfly, Bicyclus anynana, exhibits seasonal polyphenism. The wet season form has large eyespots and a pale band while these characters are much less conspicuous or absent in the dry season form. This plasticity is induced in the laboratory by use of a standard series of constant temperatures in the larval stage yielding a continuous norm of reaction. Butterflies in this study were reared from hatchling larvae in seven regimes which differed with respect to thermoperiod or photoperiod. The effect of rearing treatment on the phenotypic plasticity of the adult wing pattern, on life history traits and on larval feeding rhythms was investigated. Photoperiod had little effect except that constant light produced a higher mortality and tended to produce a longer development time. Thermoperiod had a major effect on the life history traits in comparison to a constant temperature regime with the same daily mean: development time was shorter with higher growth rates. The faster development was associated with a substantial shift in the wing pattern towards the wet season form. Larvae feed mostly at night both under constant and thermoperiod (cool nights) conditions. The results are discussed with respect to the necessity of matching field and laboratory environments in studies of norms of reaction or of life history traits where the adaptive significance of the variation is important. Fluctuating conditions in nature, especially with respect to thermoperiod, must be taken into account.     

Relationships between food resources, foraging patterns, and reproductive success in the water pipit, Anthus sp. spinoletta

A basic but rarely tested assumption in optimal foraging theoryis that positive relationships exist between the foraging patternof an animal, its short-term benefits in feeding, and its long-termfitness. We present evidence for these relationships for a centralplace foraging situation. We studied the foraging behavior ofadult water pipits (Anthus sp. spinoletta) feeding nestlingsin an Alpine habitat near Davos, Switzerland, with the followingresults: (1) searching effort decreases with increasing distancefrom the nest, (2) the amount of prey and the proportion oflarge items brought to the nest increases with increasing foragingdistance, (3) water pipits do not forage according to habitatavailability, but prefer vegetation types with the highest fooddensity (mainly grass and herbs) and avoid those with the lowest,and (4) this selectivity is only expressed when the birds foragemore than 50 m from the nest, i.e., usually outside the territory.Among the several potential interpretations of these results,the most parsimonious is that foraging decisions are based onprofitability, i.e., on the net energy gain per time unit. Additionally,we found that food conditions translate into fitness: the numberof fledglings per nest is related positively to the averageprey biomass at the foraging place and negatively to the averagedistance between the foraging place and the nest. Maximum economicdistances, which were predicted from this food-fitness relationship,agreed well with the actual foraging distances observed. Thissuggests a dose connection between foraging decisions and fitness.In addition to the theoretical issues, some conservation issuesare also briefly discussed.     

Energetic constraints and foraging efficiency

Previous research considers foraging options that differ interms of their gross rate of gain b and rate of energy expenditurec. This research argues that maximizing efficiency b/c willmaximize net energetic gain when there is an upper limit onthe amount of energy that can be assimilated. This analysisdoes not include the expenditure during the time for which theanimal is unable to forage because of this constraint. Whenthis expenditure is included, maximizing efficiency is no longeroptimal. Instead the best feeding option is the one with thehighest value of b/(c – c₁), where c, is the metabolicrate when the animal is not foraging.     

The evolution of alternative mating strategies in variable environments

Summary We assessed the influence of phenotypic plasticity in age at maturity on the maintenance of alternative mating strategies in male Atlantic salmon,Salmo salar. We calculated the fitness,r, associated with the parr and the anadromous strategies, using age-specific survival data from the field and strategy-specific fertilization data from the laboratory. The fitness of each strategy depended largely on mate competition (numbers of parr per female, i.e. parr frequency) and on age at maturity. Fitness declined with increasing numbers of parr per female with equilibrium frequencies (at which the fitnesses of each strategy are equal) being within the range observed in the wild. Equilibrium parr frequencies declined with decreasing growth rate and increasing age at maturity. Within populations, the existence of multiple age-specific sets of fitness functions suggests that the fitnesses of alternative strategies are best represented as multidimensional surfaces. The points of intersection of these surfaces, whose boundaries encompass natural variation in age at maturity and mate competition, define an evolutionarily stable continuum (ESC) of strategy frequencies along which the fitnesses associated with each strategy are equal. We propose a simple model that incorporates polygenic thresholds of a largely environmentally-controlled trait (age at maturity) to provide a mechanism by which an ESC can be maintained within a population. An indirect test provides support for the prediction that growth-rate thresholds for parr maturation exist and are maintained by stabilizing selection. Evolutionarily stable continua, maintained by negative frequency-dependent selection on threshold traits, provide a theoretical basis for understanding how alternative life histories can evolve in variable environments.     

GENETIC AND MORPHOLOGICAL DIVERGENCE AMONG MORPHOTYPES OF THE ISOPOD EXCIROLANA ON THE TWO SIDES OF THE ISTHMUS OF PANAMA

Excirolana braziliensis is a dioecious marine isopod that lives in the high intertidal zone of sandy beaches on both sides of Central and South America. It possesses no larval stage and has only limited means of adult dispersal. Indirect estimates of gene flow have indicated that populations from each beach exchange less than one propagule per generation. Multivariate morphometrics have discovered three morphs of this species in Panama, two of them closely related and found on opposite sides of Central America (“C morph” in the Caribbean and “C′ morph” in the eastern Pacific), the third found predominantly in the eastern Pacific (“P morph”). Though the P and C′ morphs are seldom found on the same beach, they have overlapping latitudinal ranges in the eastern Pacific. A related species, Excirolana chamensis, has been described from the Pacific coast of Panama. Each beach contains populations that remain morphologically and genetically stable, but a single drastic change in both isozymes and morphology has been documented. We studied isozymes and multivariate morphology of 10 populations of E. braziliensis and of one population of E. chamensis. Our objective was to assess the degree of genetic and morphological variation, the correlation of divergence on these two levels of integration, the phylogenetic relationships between morphs, and the possible contributions of low vagility, low gene flow, and occasional extinction and recolonization to the genetic structuring of populations. Genetic distance between the P morph, on one hand, and the other two morphotypes of E. braziliensis, on the other, was as high as the distance between E. braziliensis and E. chamensis. Several lines of evidence agree that E. chamensis and the P morph had diverged from other morphs of E. braziliensis before the rise of the Panama Isthmus separated the C and C′ forms, and that the P morph constitutes a different species. A high degree of genetic differentiation also exists between populations of the same morph. On the isozyme level, every population can be differentiated from every other on the basis of at least one diagnostically different locus, regardless of geographical distance or morphological affiliation. Morphological and genetic distances between populations are highly correlated. However, despite the high degree of local variation, evolution of E. braziliensis as a whole has not been particularly rapid; divergence between the C and C′ morphs isolated for 3 million yr by the Isthmus of Panama is not high by the standard of within-morph differentiation or by comparison with other organisms similarly separated. Alleles that are common in one population may be absent from another of the same morph, yet they appear in a different morph in a separate ocean. The high degree of local differentiation, the exclusive occupation of a beach by one genotype with rare arrival of foreign individuals that cannot interbreed freely with the residents, the genetic stability of populations with infrequent complete replacement by another genetic population, and the sharing by morphs of polymorphisms that are not shared by local populations, all suggest a mode of evolution concentrated in rare episodes of extinction and recolonization, possibly coupled with exceptional events of gene flow that help preserve ancestral variability in both oceans.     

SEXUAL SELECTION IN THE BARN SWALLOW (HIRUNDO RUSTICA). IV. PATTERNS OF FLUCTUATING ASYMMETRY AND SELECTION AGAINST ASYMMETRY

The patterns of variation in fluctuating asymmetry were studied in four morphological characters of the barn swallow Hirundo rustica. The level of absolute and relative asymmetry was larger in the secondary sexual character “outer tail length” than in three nonsexual morphological traits (wing, central tail, and tarsus length). The extent of individual asymmetry in outer tail length was negatively correlated with tail-ornament size, whereas the relationship between asymmetry of all other morphological characters and their size was flat or U-shaped. Asymmetry in outer tail length was unrelated to asymmetry in other morphological characters, whereas asymmetries in the length of wing, central tail, and tarsus were positively correlated. Male bam swallows exhibited larger asymmetry in outer tail length than females. Asymmetry of most morphological traits exhibited intermediate repeatabilities between years, with the exception of male and female outer tail length, which were highly repeatable. Tail asymmetry of offspring weakly, though significantly, resembled that of their parents. Asymmetry in wing and outer tail length was also significantly related to several fitness components. Male barn swallows that acquired a mate were less asymmetric in wing and outer tail length than unmated males. Females with more asymmetrical tails laid eggs significantly later. Annual reproductive success was unrelated to fluctuating asymmetry. Male barn swallows that survived were less asymmetric in wing and outer tail length than nonsurvivors, whereas female survivors were less asymmetric in outer tail length than nonsurvivors. These results suggest that levels of fluctuating asymmetry in barn swallows are associated with differences in fitness.     

Field study on the foraging characteristics of a ponerine ant,Hagensia havilandi Forel

Summary A field study of the foraging strategy used by the ponerine ant,Hagensia havilandi is reported. They have permanent nests in the leaf litter of coastal forests.H. havilandi is a diurnal forager and collects a variety of live and dead arthropods. These predatory ants exhibit individual foraging with no cooperation in the search for or retrieval of food items. Three colonies were observed and showed similar temporal and spatial foraging patterns. The paths of individual ants were followed and the results showed that the foragers exhibit area fidelity, and return to the nest via a direct route on finding on prey item. Several foragers did not return to the nest at dusk but returned the following morning. Occasionally a limited amount of tandem recruitment was displayed.     

PHYLOGENETIC RELATIONSHIPS WITHIN THE COLONIAL VOLVOCALES (CHLOROPHYTA) INFERRED FROM CLADISTIC ANALYSIS BASED ON MORPHOLOGICAL DATA1

A cladistic analysis was used to deduce the phylogenetic relationships within the colonial Volvocales. Forty-one pairs of characters related to gross morphology and ultrastructure of vegetative colonies as well as asexual and sexual reproduction were analyzed based on parsimony, using the PAUP 3.0 computer program, for 25 species belonging to nine volvocacean and goniacean genera of the colonial Volvocales. Chlamydomonas reinhardtii Dangeard was the outgroup. The strict consensus tree indicated the presence of two monophyletic groups, one composed of all the volvocacean species analyzed in this study and the other containing the goniacean species except for the four-celled species Gonium sociale (Dujardin) Warming. In addition, these two groups constitute a large monophyletic group, to which G. sociale is a sister group. A new combination Tetrabaena socialis (Dujardin) Nozaki et Itoh and a new family Tetrabaenaceae Nozaki et Itoh are thus proposed for G. sociale. In addition, the analysis suggests that the volvocacean genera Eudorina and Pleodorina are paraphyletic groups, respectively, and that the monotypic genus Yamagishiella has no autapomorphic characters and represents primitive features of the anisogamous and oogamous genera of the Volvocaceae. Phylogenetic relationships within the Volvocaceae and the Goniaceae, as well as the various modes of sexual reproduction exhibited by these organisms, are discussed on the basis of the analysis.     

A simple classification of the complex parts of vascular plants

Four operational morphological units are defined to characterize 'typical' stems, leaves, roots and hairS. Any complex part can be classified among these OMUs by a graphical representation or by a quadruplet suitable for computer-aided analyses. Thirty-nine examples are given, suggesting a continuous view of plant morphology.