Brain size, development and metabolism in birds and mammals

Recent hypotheses that variation in brain size among birds and mammals result from differences in metabolic allocation during ontogeny are tested.
Indices of embryonic and post-embryonic brain growth are defined. Precocial birds and mammals have high embryonic brain growth indices which are compensated for by low post-embryonic indices (with the exception of Homo supiens ). In contrast, altricial birds and mammals have low embryonic and high post-embryonic indices. Altricial birds have relatively small brains at hatching and develop relatively large brains as adults, but among mammals there is no equivalent correlation between variation in adult relative brain sizes and state of neonatal development.
Compensatory brain development in both birds and mammals is associated with compensatory parental metabolic allocation. In comparison with altricial development, precocial development is characterized by higher levels of brain growth and parental metabolic allocation prior to hatching or birth and lower levels subsequently. Differences between degrees of postnatal investment by the parents in the young of precocial birds versus precocial mammals may result in the different patterns of adult brain size associated with precociality versus altriciality in the two groups.
The allometric exponent scaling brain on body size differs among taxonomic levels in birds. The exponent is higher for some parts of the brain than others, irrespective of taxonomic level. Unlike mammals, the exponents for birds do not show a general increase with taxonomic level. These pattcrns call into question recent interpretations of the allometric exponent in birds. and the reason for changes in exponent with taxonomic level.     

On a polyploid complex of freshwater snails (Planorbidae: Bulinus) in Ethiopia

Allometric growth of the postembryonic stages of Ectobius lapponicus (Linn.) and E. panzeri Stephens is described and analysed. Alternative methods of computing the allometric growth equation are discussed and applied, with the necessary significance tests, to a large number of skeletal structures in all developmental stages of each sex. The validity of Dyar's Law and its modifications are assessed quantitatively.  

Summary:

The growth in linear dimensions of 74 exosketetal structures has been studied quantitatively in all instars of both sexes of Ectobius lapponicus and E. panzeri.
Almost all the investigated structures show some statistically significant degree of allometric growth, but relatively few of them grow by simple allometry. Some possible reasons for this are indicated.
Well-defined growth gradients exist in the appendages and along the main axis of the body. Allometric growth contours, showing sexual and specific differences, express the temporal and spatial variations in the allometric growth ratio.
The limitations of Dyar's Law and Przibram's Rule are discussed and it is shown that conformity with Dyar's Law is not improved by allowing for differences in the duration of the instars.
It is emphasized that allometric growth studies should involve the choice of a suitable reference dimension and the selection of an appropriate statistical model to which the growth data are fitted. The estimation of the allometry parameters a and b , as well as the appropriate significance tests, depend on the model chosen, though the two models compared in this study do not yield appreciably different estimates.     

Allometric study of the lachrymal and Harderian glands of the rat during postnatal life.

The authors performed an allometric study of the growth of the rat's lachrymal and Harderian glands, during postnatal life. From the analysis of the results, they could conclude: (1) the growth of these glands in relation to body weight, during postnatal life, could be considered similar, following the allometric law; (2) the differential growth of the glands occurred in two stages: from birth until the 15th day and from the 15th day until the final period of life studied; (3) the two stages of development were separated by a critical period, during which an abrupt modification of the allometric coefficient occurred; (4) during the first days of postnatal life, the development of the Harderian gland was characterized by a high rate of growth and, just after eyelid disjunction and during rest of postnatal life, by a rate of allometric of growth less than 1. It is interesting to observe that the lachrymal and Harderian glands' critical period of development on the 15th day of postnatal life coincides with the time at which the eyelids of the animal open.     

Analytical morphometry of sagittal fronto-facial profiles of 2nd-6th month human foetus

A prominent problem in the morphological study of developing biological structures is shape parametrization with the aim to evaluate transformations in a non subjective way. We carried out an analytical morphometrical study on a series of human embryos, by means of the S.A.M. (Shape Analytical Morphometry) software package. After standardization and normalization of the fronto-facial sagittal profiles, the analytical procedures applied were: 1) Fourier analysis: each profile was considered as an irregular but periodic function obtained by the sum of sinusoids of increasing order. 2) Shape Asymmetry Evaluation: couples of profiles are compared by means of "Janus" procedure; by a parabolic fitting we obtained parameters able to evaluate allometric and isometric differences between the two profiles. The preliminary results of the applied procedures indicated that rate and direction of allometric and isometric growth are not constant during the time.     

Do Rotifer Jaws Grow After Hatching?

The hard articulated jaws of some pseudocoelomate metazoans were recently used in reconstructing their phylogenetic relationships, but we still do not know if these structures could change in size and shape during the life of individuals, and experimental data are lacking on their post-embryonic development. Rotifers are one of the groups in which hard articulated jaws, called trophi, are well known, and are widely used taxonomically. Here we report on SEM study of trophi of rotifers of different ages, to determine if the trophi structures change in shape and/or in size during post-embryonic development. We used linear measurements and geometric morphometrics analyses from scanning electron microscopic pictures of trophi of Cupelopagis vorax, Dicranophorus forcipatus, Macrotrachela quadricornifera, Notommata glyphura, Rotaria macrura, R. neptunoida, and R. tardigrada. Results for these species show that trophi do not change after hatching, either in size or in shape. In contrast, data on Asplanchna priodonta reveal trophi growth after hatching.     

Post-embryonic development of amphipod crustacean pleopods and the patterning of arthropod limbs

Morphology and post-embryonic development of the pleopodal rami of the amphipod crustacean Gammarus roeselii Gervais, 1835 are described, with particular reference to the differentiation of limb articles. An article is the part of a limb between two complete arthrodial membranes and, in the pleopodal rami, most arthrodial membranes are accompanied by two plumose setae on the opposite faces of the article. During post-embryonic development, the rami acquire new plumose setae and new arthrodial membranes proximal to the already differentiated part; this implies that new articles are produced by the division of the most proximal article only. On the proximal article of the inner ramus there are also some bifid setae, whose number increases post-embryonically, most likely by proximal addition of new setae. Data from different limb types of different arthropod groups suggest that during post-embryonic development new articles are also produced from one or more specific zone(s) of the limbs. General aspects of these “growth zones” in arthropod limbs are discussed and their distribution, in respect to both different limb types and different taxa, is reviewed. The available data do not allow unambiguous conclusions about the evolution of growth zones and limb patterning, but the most relevant questions are nevertheless outlined.     

Abstracts of papers to be presented at the Fiftieth Annual Meeting of the American Association of physical anthropologists Detroit,Michigan April 22–25, 1981

The critical problem confronting all allometric studies is the choice of an appropriate size variable, especially when body mass or some other measure of total size is unavailable. A method proposed by Jolicoeur (1963a,b) claims to generate an internal size variable by a principal components analysis of the covariance matrix of logarithmically transformed data, from which allometric coefficients can be computed. Despite the current popularity of this method, the precise relationship and degree of compatibility between such multivariate coefficients and the exponent of the bivariate power function (Y = βX^α) is unknown. This study evaluates the comparability and interpretability of allometric values computed by Jolicoeur's procedure and by standard bivariate regressions (least squares and major axis). Two primate data sets with known measures of size were utilized for these purposes: (1) longitudinal growth data from radiographs of two species of capuchin monkeys, Cebus apella and Cebus albifrons; and (2) interspecific osteometric data from a series of adult lemurs, species of prosimians from Madagascar. Consistent differences exist between multivariate and bivariate allometric coefficients for both ontogenetic and static data sets. Multivariate analysis underestimated the coefficients in the former and overestimated them in the latter. The internal size variable generated by principal components analysis is clearly not equivalent to, and hence not a suitable substitute for, known measures of size. Moreover, multivariate coefficients are very sensitive to the composition of variables in a given data set; α values of a variable changed appreciably depending on the other variables included in the analysis. The multivariate coefficients are also sample-specific, and provide misleading biological information when compared between samples (e.g., between species of capuchin monkey). For allometric investigations designed to evaluate scaling parameters relative to total size, alternative analytical solutions to the Jolicoeur method should be considered.     

Homoplasy and the evolution of ontogeny in papionin primates

Recent advances in developmental biology reveal that patterns of morphological development, even during early phases, may be highly susceptible to evolutionary change. Consequently, developmental data may be uninformative with regard to distinguishing homology and homoplasy. The present analysis evaluates postnatal ontogeny in papionin primates to test hypotheses about homology and homoplasy during later periods of development. Specifically, the analysis studies the allometric bases of craniometric resemblances among four papionin genera to test the hypothesis that homoplasy in adult cranial form, particularly of baboons (Papio) and mandrills (Mandrillus), is underwritten by divergent patterns of development. Bivariate and multivariate allometric analyses demonstrate that the developmental patterns in Papio baboons diverge markedly from ontogenetic allometric trajectories in other papionin species. The resemblances between Papio and Mandrillus (assuming that patterns of development in smaller papionins are ancestral) are largely consequences of perinatal increases in relative brain size in juvenile Papio. Postnatal growth to large size and strong negative allometry of neurocranial form results in shape similarities because developmental pathways for large papionin genera intersect. Analyses show that allometric data may not be particularly informative in revealing homoplasy. However, placed into proper phylogenetic context, such data illustrate derived patterns of development that may reflect critically important life-history or ontogenetic adaptations.     

Plant Growth Analysis: Allometry, Growth and Interference in Orchardgrass and Timothy

A multiple regression procedure was used to evaluate allometricresponses to stand age and species population densities in monoculturesand mixtures of orchardgrass (Dactylis glomerata L., also knownas cocksfoot) and timothy (Phleum pratense L.). In each speciesthe allometry between shoot dry weight and either leaf areaor tiller number per plant was studied. Population density treatmentsaffected allometry by changing allometric exponents expressingthe ratio of relative growth rates of different plant characteristics.Allometric relationships changed as growth proceeded, and thetwo species differed in their allometric responses to treatments. Plant growth analysis, allometry, competition, Dactylis glomerata L., Phleum pratense L.     

How mantids gain insight into the new maximum catching distance after each ecdysis

The maximum catching distance (MCD) of which a mantid, Stagmatoptera biocellata, can successfully strike a prey keeps a certain relationship with its foreleg extension in each instar. Two hypotheses aimed at explaining the occurrence of this relationship in spite of the sudden growth in dimensions that takes place after each ecdysis are discussed. The assumption that structures involved in depth perception show changes that allow the mantid to estimate correctly the MCD is tested. This ‘automatic adaptation’ hypothesis receives strong support from the results. They indicate that throughout postembryonic development the MCD has a constant proportionality with three head dimensions, i.e. the head breadth, the ocular globe breadth, and the ocular prominence. These modifications during the saltatory growth cause a change in the optical triangulation system that can, by itself, explain the perfect estimate of the distance within which a strike is a worth-while effort, without resorting to a learning hypothesis.     

Allometric growth and adaptive value of exaggerated body parts in the larvae of Gratiana spadicea (Klug) (Coleoptera: Chrysomelidae)

1. Larvae of tortoise beetles present exaggerated body parts in association with an abdominal shield, which is made of faeces and exuviae that are deposited on the urogomphi throughout ontogeny. Growth trajectories and scaling relationships of these functional structures associated with the shield, if any, are unknown. 2. This study of Gratiana spadicea first tested, under field conditions, whether there is adaptive value associated with the shield regarding protection against predation and sunlight. Then, under laboratory conditions, the growth trajectory and allometric relationships among body parts were investigated, including scoli, individual and apparent furcae, and shield. The influence of food deprivation on the development of these structures was also determined. 3. Findings from previous studies were confirmed, suggesting that the adaptive value assigned to the shield is related to protection against predators. The present study demonstrated for the first time that the shield acts as a parasol in cassidines, decreasing the exposure of their larval body to sunlight. The scoli and apparent furca are exaggerated structures of G. spadicea, the development of which involves allometric growth and greater energetic investment (positive allometry) during ontogeny. There was proportionally less energetic investment for somatic construction of individual furca (negative allometry) due to the accumulation of the exuviae. 4. The possible consequences, in terms of developmental costs and survivorship benefits associated with the evolution of such exaggerated structures, are discussed.     

The nature and basis of sexual dimorphism in the primate skeleton

This study sets out to document and analyse sexual dimorphism in the teeth and bones of five primate groups, including man. Specimens were only included in the analysis if their sexual attribution was reliable and was based on non-osteological criteria. Ninety raw measurements, both cranial and post-cranial, were used and 11 indices were computed from them.
The parameters of each sample were computed in order to compare these results with previous estimates of dimorphism which have not always been based on reliably sexed samples. Correlation matrices were computed.
The overall sex differences were subdivided into "shape" and "size" components using Penrose's size and shape distances and by computing the principal components of each data set. The visually apparent shape differences were confirmed metrically and then examined to see whether they could be explained by allometric effects or whether there was evidence for sexual differences in growth patterns.
Using femur length as the independent variable, because of its correlation with overall size, allometric coefficients were computed for the logarithmically transformed data. The coefficients were in some cases very different between sexes but the majority did not achieve statistical significance. Of those that were significantly different only in Homo did such differences narrowly exceed the number that would be expected by chance alone.
As the vast majority of allometric coefficients for the pooled male and female data differed from unity, the hypothesis that most of the considerable shape differences that exist between some male and female primates are due to underlying growth differences must be rejected. It is suggested that such differences are simply the result of disproportionate change in size.     

Evolution of ontogeny in the hippopotamus skull: using allometry to dissect developmental change

Allometry describes the effect of size change on aspects of an organism's form and can be used to summarize the developmental history of growing parts of an animal. By comparing how allometric growth differs between species, it is possible to reveal differences in their pathways of development. The ability to compare and categorize developmental change between species is demonstrated here using morphometric methods. This involves the interspecific statistical comparison of a large number of bivariate relationships that summarize ontogenetic trajectories. These linear ontogenetic trajectories can be modified as they evolve in any of three ways: ontogenetic scaling indicative of change in the duration of growth, lateral shifts indicative of changes in prenatal development, and directional change indicative of novel modes of postnatal growth. I apply this analysis to skulls of the common hippopotamus ( Hippopotamus amphibius ) and the pygmy hippopotamus ( Hexaprotodon liberiensis ). The number of allometric changes falling into each category was statistically determined and Jolicoeur's multivariate generalization of simple allometry was used to provide an overview of cranial variation. For these skulls, directional change was not found to be statistically significant, but ontogenetic scaling and lateral shifts were both common. This indicates that conserved patterns of growth covariance (ontogenetic scaling) can be separated from novel or derived patterns (directional change and/or lateral shifts). This study demonstrates that He. liberiensis is not simply an ontogenetically scaled version of its larger relative. The evolutionary implications of allometric growth variation are discussed in the light of these findings and those of other studies.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 625–638.     

Egg mass and incubation period allometry in birds and reptiles: effects of phylogeny

Here we test the hypothesis that the relationship between egg mass at oviposition (IEM) and incubation period ( I _p) is a function of the taxonomic relatedness of bird and reptile species. Allometric relationships between IEM and I _p were examined for 1525 bird species and 201 reptilian species. Treating species as independent data revealed the allometric exponent linking I _p to IEM to be 0.234 for birds and 0.138 for reptiles. However, ANCOVA revealed that within both birds and reptiles the elevation and slope of the regression lines were dependent on the taxonomic order studied, indicating that the exponents were confounded by the phylogenetic relatedness of species. Thus, allometric exponents were recalculated based on the method of comparative analysis using independent contrasts. This technique revealed that the allometric exponent in both birds and reptiles was confounded by phylogeny. In birds the allometric relationship between I _p and IEM was almost halved to 0.122, whereas in reptiles the exponent increased to 0.185. Importantly, the results demonstrate that some results of allometric analyses can be artefacts of the method of analysis of the dataset. That for bird eggs I _p is not determined in large part by egg mass allows new questions to be posed regarding the ecological and physiological factors affecting the length of incubation, and hence rates of embryonic growth, for different taxa and habitats.     

从弹尾纲和原尾纲的亲缘关系质疑缺尾纲（=近昆虫纲）的有效性

鉴于弹尾目（跳虫）和原尾目的尾部都没有尾须（cerci）,Brner于1910年就把这两类归并为一个类群缺尾纲（Ellipura）,这一分类阶元长期被许多昆虫学家沿用至今。Kukalová-Peck(1987 )在讨论化石双尾虫（?）附肢的总体结构（ground plan）时,认为跳虫和原尾虫的腹部侧板更原始,附肢无转节,将二者归纳成近昆虫纲（Parainsecta）。但是从形态特征、内部结构、比较精子学、变态类型和胚后发育等的特点以及线粒体DNA和核糖体DNA的测序分析结果,显示弹尾纲与原尾纲之间存在诸多重要差异,不具备较为密切的亲缘关系,我们不支持（弹尾纲+原尾纲）组成缺尾纲或近昆虫纲。据此建议取消缺尾纲（=近昆虫纲）这一分类阶元。     

Allometry and adaptation of body proportions and stature in African pygmies

We have analyzed the growth allometry of external body proportions in Efe pygmies from Zaire and combined these data with values from the literature for comparable dimensions in adult pygmies and nonpygmies. We sequentially tested the hypotheses that adult proportion differences between 1) male vs. female Efe, and 2) pygmies vs. nonpygmies result from ontogenetic scaling, or the differential extension of common patterns of growth allometry. Results indicate an almost complete concordance of allometric trajectories for male and female Efe. These preliminary analyses also strongly suggest that adult nonpygmy Africans generally differ from pygmies in their terminal size and correlated allometric consequences, rather than in more fundamental alterations of underlying patterns of growth. Biacromial diameter emerges as the measurement most likely to depart from this general pattern. These results provide further evidence that shifts in systemic growth hormones yielding differences in terminal overall body size may be accompanied by global and coordinated allometric transformations. Certain proportion differences previously interpreted by some as specific evidence of primitive retention in pygmies in fact reflect simple growth allometric correlates of the derived rapid size decrease in these groups. Selected divergent body proportions characterizing adult pygmies, previously interpreted by some as independent evidence of climatic adaptation, also reflect such allometric correlates of ontogenetic scaling. We critically assess arguments that the small overall body size of pygmies was specifically selected for reasons of thermoregulatory efficiency, and consider an alternative or complementary scenario, based on selection for small size in order to reduce caloric requirements. © 1996 Wiley-Liss, Inc.     

Phenotypic analysis of adults and eggs of Fasciola hepatica by computer image analysis system

Knowledge of the morphological phenotypes of the liver fluke Fasciola hepatica (Trematoda: Digenea) is analysed. The influence of parasite age on its dimensions, the adult fluke growth model, variation in a biometric variable versus time, and variation in a biometric variable versus another biometric variable (allometric model) are revised. The most useful allometric model appears to be (y2 m-y2)/y2=c [(y1 m-y1)/y1](b), where y1=body area or body length, y2=one of the measurements analysed, y1 m, y2 m=maximum values towards which y1 and y2, respectively, tend, and c, b=constants. A method based on material standardization, the measurement proposal and allometric analysis is detailed. A computer image analysis system (CIAS), which includes a colour video-camera connected to a stereomicroscope (for adult studies) and a microscope (for egg studies), facilitates the processing of digital imaging. Examples of its application for the analysis of the influence of different factors on the liver fluke phenotype are shown using material from the Northern Bolivian Altiplano, where human and domestic animal fascioliasis is caused by F. hepatica only. Comparisons between the development of livestock fluke populations from highlands and lowlands are discussed and the relationships between host species and liver fluke morphometric patterns is analysed.     

Positional specification in the segmental growth pattern of an early arthropod

In many arthropods, there is a change in relative segment size during post-embryonic development, but how segment differential growth is produced is little known. A new dataset of the highest quality specimens of the 429 Myr old trilobite Aulacopleura koninckii provides an unparalleled opportunity to investigate segment growth dynamics and its control in an early arthropod. Morphometric analysis across nine post-embryonic stages revealed a growth gradient in the trunk of A. koninckii. We contrastively tested different growth models referable to two distinct hypotheses of growth control for the developing trunk: (i) a segment-specific control, with individual segments having differential autonomous growth progression, and (ii) a regional control, with segment growth depending on their relative position along the main axis. We show that the trunk growth pattern of A. koninckii was consistent with a regional growth control producing a continuous growth gradient that was stable across all developmental stages investigated. The specific posterior-to-anterior decaying shape of the growth gradient suggests it deriving from the linear transduction of a graded signal, similar to those commonly provided by morphogens. A growth control depending on a form of positional specification, possibly realized through the linear interpretation of a graded signal, may represent the primitive condition for arthropod differential growth along the main body axis, from which the diverse and generally more complex forms of growth control in subsequent arthropods have evolved.     

The effects of reduced and oxidized glutathione on white spruce somatic embryogenesis

Summary The glutathione-glutathione disulfide redox pair was utilized to improve white spurce somatic embryo development. Mature cotyledonary-stage somatic embryos were divided into two groups (A and B) based on morphological normality and the ability of the mature somatic embryos to convert into plantlets. Group A embryos had four or more cotyledons and converted readily upon germination after a partial drying treatment. Group B embryos had three or fewer cotyledons with a low conversion frequency. The addition of reduced glutathione (GSH) at a concentration of 0.1 mM resulted in an increase in embryo production (total population) with a mean total number of 64 embryos per 100 mg embryogenic tissue as well as an increase in post-embryonic root growth. However, at a higher concentration (1 mM), GSH inhibited embryo formation. The manipulation of the tissue culture environment via the inclusion of glutathione disulfide (GSSG), at concentrations of 0.1 and 1.0 mM, enhanced the development of better-quality embryos. This quality was best exemplified when embryos forming four or more cotyledons increased by at least twofold to 73.9% when treated with 1.0 mM GSSG, compared to 38% in control. Furthermore, this improved quality was reflected by an increased conversion frequency. A 20% increase in the ability of the somatic embryo to produce both root and shoot structures during post-embryonic development was noted when embryos were matured on maturation medium supplemented with 1.0 mM GSSG over the control.