The roles of cell size and cell number in determining ovariole number in Drosophila

All insect ovaries are composed of functional units called ovarioles, which contain sequentially developing egg chambers. The number of ovarioles varies between and within species. Ovariole number is an important determinant of fecundity and thus affects individual fitness. Although Drosophila oogenesis has been intensively studied, the genetic and cellular basis for determination of ovariole number remains unknown. Ovariole formation begins during larval development with the morphogenesis of terminal filament cells (TFCs) into stacks called terminal filaments (TFs). We induced changes in ovariole number in Drosophila melanogaster by genetically altering cell size and cell number in the TFC population, and analyzed TF morphogenesis in these ovaries to understand the cellular basis for the changes in ovariole number. Increasing TFC size contributed to higher ovariole number by increasing TF number. Similarly, increasing total TFC number led to higher ovariole number via an increase in TF number. By analyzing ovarian morphogenesis in another Drosophila species we showed that TFC number regulation is a target of evolutionary change that affects ovariole number. In contrast, temperature-dependent plasticity in ovariole number was due to changes in cell-cell sorting during TF morphogenesis, rather than changes in cell size or cell number. We have thus identified two distinct developmental processes that regulate ovariole number: establishment of total TFC number, and TFC sorting during TF morphogenesis. Our data suggest that the genetic changes underlying species-specific ovariole number may alter the total number of TFCs available to contribute to TF formation. This work provides for the first time specific and quantitative developmental tools to investigate the evolution of a highly conserved reproductive structure.     

Comparative fecundity, clutch size, ovariole number and egg size of Dacus tryoni and D. jarvisi, and their relationship to body size

Dacus tryoni (Frogg.) (Diptera: Tephritidae) is the main tephritid pest of cultivated fruits in Australia. D. jarvisi (Tryon) is also able to infest these fruits. Some factors influencing the rate at which D. tryoni and D. jarvisi exploit patches of host fruits were examined to determine whether one species may have an advantage when they exploit the same fruits in the field. Measurements of the intrinsic rate of increase, ovariole number, clutch size and egg size and the influence of body size on these parameters were made for both species in the laboratory. Up to 10 weeks of age D. tryoni produced twice as many eggs as D. jarvisi, most during a peak 3–5 weeks after adult emergence. The difference in fecundity up to 10 weeks can be explained partly by the higher number of ovarioles in D. tryoni (38/ovary) compared to D. jarvisi (27/ovary). In addition D. tryoni produces smaller eggs than D. jarvisi and distributes them in smaller clutches; 3–4 eggs/clutch vs 10–15. In both species there was a positive correlation between ovariole number and body size (as measured by wing length). By contrast, egg size remained constant over a broad range of body sizes. The influence of these life history differences on the interaction between D. tryoni and D. jarvisi in the field is discussed.

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Résumé D. tryoni Frogg est la principale téphrididae nuisible aux fruits cultivés en Australie. Cependant, plusieurs autres espèces de Dacus, dont D. jarvisi Tryon sont susceptibles de contaminer ces fruits. Quelques facteurs influant sur le taux de contamination de fruits ont été examinés pour déterminer si une espèce peut être avantagée lors de l'exploitation des mêmes fruits dans la nature. L'influence de la taille de l'adulte sur le taux d'accroissement intrinsèque, sur le nombre d'ovarioles, sur l'effectif des pontes et sur la taille des ufs a été examinée au laboratoire sur les 2 espèces. Pendant les 10 premières semaines, D. tryoni produit 2 fois plus d'ufs que D. jarvisi, la plupart étant pondus entre la 3^e et la 5^e semaines après l'émergence. La production de D. tryoni diminue rapidement après ce maximum. D. jarvisi ne présente pas ce maximum précoce, et la production des ufs se fait au même rythme entre les 3^e et 7^e semaines, avant de diminuer graduellement. La différence de fécondité au bout de 10 semaines peut être expliquée partiellement par le plus grand nombre d'ovarioles de D. tryoni (38/ovaire) contre 24/ovaire pour D. jarvisi. De plus, D. tryoni forme des ufs plus petits que D. jarvisi et l'effectif de chacune de ses pontes est plus limité: 3 à 4 ufs contre 10 à 15. Chez les deux espèces, il y a une relation directe nette entre le nombre d'ovarioles et la taille du corps de la femelle (mesurée par la longueur de l'aile). Par contre, la taille des ufs est indépendante d'une grande gamme de tailles du corps des femelles. Les caractéristiques biologiques de D. tryoni le rendent capable de contaminer rapidement les bouquets de fruits qu'il vient de coloniser, ce qui réduit les disponibilités pour les autres espèces susceptibles de contaminer ces fruits. Dans le cas particulier de D. jarvisi, D. tryoni a peu d'impact sur son niveau de population, puisque D. jarvisi peut exploiter aussi son hôte d'origine, Planchonia careya: Il ne tend à entrer en compétition avec D. tryoni que pour une ou deux générations tardives en été, quand l'hôte partage (la goyave) est souvent abondant. Néanmoins, si les fruits sont rares à cette époque ou si les 2 espèces sont obligées de partager leurs hôtes pendant plusieurs générations (hors de l'aire de Planchonia), D. tryoni aura un avantage certain.

     

Relationship between collagen fibril diameters and body size

Summary Suspensions of collagen fibrils obtained from derma of Elasmobranchia and Actinopterygia of different body sizes and developmental stages were examined by transmission electron microscopy. Fibril diameters were measured and classified into groups comprising a 20 nm diameter interval. Diagrams showing fibril populations of each fish were made. The measurements were averaged and their confidence intervals and standard errors determined. For each species other diagrams were plotted in which the mean diameters were correlated to the body length of each sample. The results show that: 1) a correlation exists between an increase in diameter of collagen fibrils and somatic growth until sexual maturity is reached; 2) fibril populations are subsequently spread over a wider range due to the presence in the derma of classes of newly formed and therefore thinner fibrils. The deposition of new fibrils is possibly influenced by individual factors; 3) no relationship exists between mean fibril diameter and body size; 4) no relationship exists between phylogenetic position and pattern of diameter distribution.Research supported by a grant from C.N.R. Roma (69.02087.0115.1150)     

Relationship between evenness and body size in species rich assemblages

Evenness is a key measure of community structure. Here, we examine the relationship between evenness and size–abundance distributions for both individuals and species using data gathered from Amazonian fish assemblages. We show that evenness increases as the fraction of numerically abundant species in larger body-size classes rises. As any processes that enable larger bodied species to increase their numerical dominance will influence evenness, these results help explain why evenness is an important correlate of ecosystem function.     

A test of three hypotheses for ovariole number determination in the grasshopper Romalea microptera

Ovariole number in insects determines potential fecundity and can be influenced by genes, environmental conditions during development and parental effects. In the present study, three hypotheses are tested for ovariole number determination in the grasshopper Romalea microptera (Beauvois), which exhibits both intra‐ and interpopulation variation in ovariole number. In hypothesis 1, variation in ovariole number is a result of genetic variation. In hypothesis 2, ovariole number is influenced by nutrition during development. In hypothesis 3, ovariole number is influenced by maternal nutritional status. Females from four treatments are compared: low‐food, high‐food, daughters of low‐food, and daughters of high‐food. There is a relationship between parent and offspring ovariole number despite different environments, supporting hypothesis 1. Also, ovariole numbers are slightly, but significantly lower in individuals fed a low‐food diet compared with a high‐food diet, supporting hypothesis 2. Hypothesis 3 is not supported: starved and well‐fed females produce eggs of similar mass, as well as offspring with similar numbers of ovarioles, suggesting that the nutritional status of mothers does not influence offspring mass or offspring ovariole number. The results imply that genetic variation and developmental conditions determine ovariole number in this species but maternal environment does not. These results conflict with previous studies of ovariole determination in grasshoppers and locusts.     

Relationship between adiposity and body size reveals limitations of BMI

The aims of this study were to assess 1) whether the stature-adjusted body mass index (BMI) is a valid proxy for adiposity across both athletic and nonathletic populations, and 2) whether skinfold measurements increase in proportion to body size, thus obeying the principle of geometric similarity. The research design was cross-sectional, allowing the relationship between skinfold calliper readings (at eight sites and between specific athletic and nonathletic groups, n = 478) and body size (either mass, stature, or both) to be explored both collectively, using proportional allometric MANCOVA, and individually (for each site) with follow-up ANCOVAs. Skinfolds increase at a much greater rate relative to body mass than that assumed by geometric similarity, but taller subjects had less rather than more adiposity, calling into question the use of the traditional skinfold-stature adjustment, 170.18/stature. The best body-size index reflective of skinfold measurements was a stature-adjusted body mass index similar to the BMI. However, sporting differences in skinfold thickness persisted, having controlled for differences in body size (approximate BMI) and age, with male strength- and speed-trained athletes having significantly lower skinfolds (32% and 23%, respectively) compared with controls. Similarly, female strength athletes had 29% lower skinfold measurements compared to controls, having controlled for body size and age. These results cast serious doubts on the validity of BMI to represent adiposity accurately and its ability to differentiate between populations. These findings suggest a more valid (less biased) assessment of fatness will be obtained using surface anthropometry such as skinfolds taken by experienced practitioners following established procedures.     

Relationship between body size and long bone lengths in Pan and Gorilla

Measurements of body length (vertex to heel) were abstracted from the field notes of Pan and Gorilla specimens from the Powell-Cotton Museum. Bicondylar femur and humerus length were measured on each skeleton and correlation coefficients with body length were computed. In both the separate sex and the combined sex samples of Gorilla, and in the combined sex sample of Pan, long bone lengths are significantly correlated with body size, but in Pan only 20% of the variance in body length is reflected in the long bone measurements.     

Relationship between subunit size and number of rare electrophoretic alleles in human enzymes

Data from published sources were used to compare the numbers of different electrophoretic alleles of 29 monomeric and dimeric human enzymes to their respective subunit molecular weights. Only those human enzymes were considered for which the total sample sizes were in excess of 2000 individuals. Correlations between these two variables were determined within sample size ranges of 2000≤n≤3000 and 4000≤n≤5000 individuals, and separately by quaternary class. There was no statistically significant correlation observed for the smaller sample size range in monomers; however, the correlations for the larger sample size range in monomers and both ranges in dimers were significant. Since there is no relationship between subunit size and heterozygosity, the relationships are due primarily to the incidence of rare alleles. These findings demonstrate the effect of locus-specific mutation rates, expected as a consequence of variation of cistron sizes, and imply that other forces are responsible for the relative frequencies of common alleles at some of the loci.     

Relationship between number of muscles,behavioral repertoire size,and encephalization in mammals

Behavior for mammals is built out of multiple muscles acting in a coordinated fashion. Prima facie, there are three principal ways to increase an animal's behavioral repertoire size. The first is to, for each new behavior type, create a set of new muscle types (e.g. triceps, sartorius, etc.) with new functions specifically devoted to the implementation of that behavior type. If this were the case, then although each behavior is built out of many muscles, behavior is not built in a combinatorial fashion out of muscles. The second is similar to the first in that new behavior types are implemented via new muscle types, but, instead, muscles are used in a combinatorial fashion, so that it is the combination of the new muscle type with existing muscle types that makes the new behavior type possible. This is analogous to the addition of new words in a language. The third way behavioral complexity may be scaled up is to increase the complexity of behavioral expressions themselves (rather than increasing the number of muscles types), namely by having more muscles involved in an average behavior. This is analogous to uttering longer sentences in a language. My main task in this paper is to examine which of these ways underlies the increase of behavioral complexity among mammals. Behavioral repertoire sizes from the ethology literature were accumulated for mammals from two dozen species across eight orders, and the number of muscle types was estimated from atlases of anatomy across eight mammalian orders. The manner in which behavioral complexity actually increases among mammals appears to favor the second possibility mentioned above: greater behavioral complexity is achieved primarily by increasing the number of muscle types, and by using muscles in a combinatorial fashion. The theoretical framework I describe allows us to interpret the manner in which the number of muscle types scales with behavioral repertoire size, and I conclude that the number of degrees of freedom in the construction of behavioral expressions is on the order of three, which is probably due to neurobiological limitations in forming behaviors. The ontogeny of behavior in rat is also discussed within this framework. Finally, I show that there is a strong positive relationship between behavioral repertoire size and encephalization among mammals.     

Variations in body melanisation,ovariole number and fecundity in highland and lowland populations of Drosophila melanogaster from the Indian subcontinent

We investigated geographical variations in three fitness‐related traits (body melanisation, ovariole number and fecundity) in laboratory‐reared offspring of 10 populations of Drosophila melanogaster. The populations were collected from adjacent lowland and highland localities (～80–100 km apart) in the tropical as well as subtropical regions (11.15–31.06 °N) covering a linear distance about 3 000 kilometers from south to north on the Indian subcontinent. Persistence of within‐as well as between‐population differences at 21 °C suggest that observed variations in fitness‐related traits have a genetic basis. Populations from higher altitudes showed consistently higher trait values (1.4‐fold increase) as compared with their corresponding lowland populations. By contrast, latitudinal variations were about two‐fold higher across the entire continent. Along latitude as well as altitude, population means showed higher correlation values (r > 0.98) between all the three fitness traits. However, on the basis of within‐population analysis (assorted darker and lighter flies), changes in body melanisation were significantly correlated with fecundity but not with ovariole number. Thus, analysis of within‐population trait variability should be preferred as compared with data on population means for adaptive significance of fitness‐related traits. In the present study, the role of climatic selection is evident from regression analysis with changes in annual average temperature of the sites of origin of populations along latitude as well as altitude.     

Variations in body melanisation,ovariole number and fecundity in highland and lowland populations of Drosophila melanogaster from the Indian subcontinent

We investigated geographical variations in three fitness-related traits （body melanisation, ovariole number and fecundity） in laboratory-reared offspring of 10 populations of Drosophila melanogaster. The populations were collected from adjacent lowland and highland localities （-80-100 km apart） in the tropical as well as subtropical regions （11.15-31.06°N） covering a linear distance about 3 000 kilometers from south to north on the Indian subcontinent. Persistence of within- as well as between-population differences at 21 ℃ suggest that observed variations in fitness-related traits have a genetic basis. Populations from higher altitudes showed consistently higher trait values （1.4-fold increase） as compared with their corresponding lowland populations. By contrast, latitudinal variations were about two-fold higher across the entire continent. Along latitude as well as altitude, population means showed higher correlation values （r 〉 0.98） between all the three fitness traits. However, on the basis of within-population analysis （assorted darker and lighter flies）, changes in body melanisation were significantly correlated with fecundity but not with ovariole number. Thus, analysis of within-population trait variability should be preferred as compared with data on population means for adaptive significance of fitness-related traits. In the present study, the role of climatic selection is evident from regression analysis with changes in annual average temperature of the sites of origin of populations along latitude as well as altitude.     

Relationship between measures of body size and composition and velocity of lactate threshold

The purpose of this investigation was to explore the relationship between velocity of lactate threshold (vLT) and various measures of body mass and composition: mass, lean mass, fat mass, percent body fat (% fat), and body surface area (BSA). We hypothesized that mass would be inversely related to vLT, and that differences in measures of body mass and composition would account for a significant amount of variability in vLT. A total of 21 healthy male runners served as subjects. Body composition was assessed by hydrostatic weighing. A significant negative relationship (r = -0.759, p < or = 0.01) was between body mass and vLT. The coefficient of determination between vLT and body mass indicated that nearly 58% of the variability in vLT was explained by body mass in these subjects. Significant relationships were also between vLT and BSA (r = -0.72, p < or = 0.01), fat mass (r = -0.70, p < or = 0.01), % fat (r = -0.59, p < or = 0.01), and lean mass (r = -0.41, p < or = 0.05). Linear regression yielded the following model: y = 369.48 - 1.7343 (X), where y = predicted vLT (m.min(-1)) and X = body mass (kilograms) (SEE = 15.45). Velocity of lactate threshold was significantly inversely related with body mass in a group of male runners. The calculated coefficient of determination suggests that nearly 58% of the variability in vLT was explained by body mass. The present data suggest consideration of categorizing participants in 'road runs' by body mass to equate competition, as is done in other sports (e.g., weightlifting).     

High-resolution quantitative trait locus mapping reveals sign epistasis controlling ovariole number between two Drosophila species

Identifying the genes underlying genetically complex traits is of fundamental importance for medicine, agriculture, and evolutionary biology. However, the level of resolution offered by traditional quantitative trait locus (QTL) mapping is usually coarse. We analyze here a trait closely related to fitness, ovariole number. Our initial interspecific mapping between Drosophila sechellia (8 ovarioles/ovary) and D. simulans (15 ovarioles/ovary) identified a major QTL on chromosome 3 and a minor QTL on chromosome 2. To refine the position of the major QTL, we selected 1038 additional recombinants in the region of interest using flanking morphological markers (selective phenotyping). This effort generated approximately one recombination event per gene and increased the mapping resolution by approximately seven times. Our study thus shows that using visible markers to select for recombinants can efficiently increase the resolution of QTL mapping. We resolved the major QTL into two epistatic QTL, QTL3a and QTL3b. QTL3a shows sign epistasis: it has opposite effects in two different genetic backgrounds, the presence vs. the absence of the QTL3b D. sechellia allele. This property of QTL3a allows us to reconstruct the probable order of fixation of the QTL alleles during evolution.     

The relationship between nephron number, kidney size and body weight in two inbred mouse strains

While some reports in humans have shown that nephron number is positively correlated with height, body weight or kidney weight, other studies have not reproduced these findings. To understand the impact of genetic and environmental variation on these relationships, we examined whether nephron number correlates with body weight, kidney planar surface area, or kidney weight in two inbred mouse strains with contrasting kidney sizes but no overt renal pathology: C3H/HeJ and C57BL/6J. C3H/HeJ mice had smaller kidneys at birth and larger kidneys by adulthood, however there was no significant difference in nephron number between the two strains. We did observe a correlation between kidney size and body weight at birth and at adulthood for both strains. However, there was no relationship between nephron number and body weight or between nephron number and kidney size. From other studies, it appears that a greater than two-fold variation is required in each of these parameters in order to demonstrate these relationships, suggesting they are highly dependent on scale. Our results are therefore not surprising since there was a less than two-fold variation in each of the parameters examined. In summary, the relationship between nephron number and body or kidney size is most likely to be demonstrated when there is greater phenotypic variation either from genetic and/or environmental factors.     

The relationship between nephron number,kidney size and body weight in two inbred mouse strains

While some reports in humans have shown that nephron number is positively correlated with height, body weight or kidney weight, other studies have not reproduced these findings. To understand the impact of genetic and environmental variation on these relationships, we examined whether nephron number correlates with body weight, kidney planar surface area, or kidney weight in two inbred mouse strains with contrasting kidney sizes but no overt renal pathology: C3H/HeJ and C57BL/6J. C3H/HeJ mice had smaller kidneys at birth and larger kidneys by adulthood, however there was no significant difference in nephron number between the two strains. We did observe a correlation between kidney size and body weight at birth and at adulthood for both strains. However, there was no relationship between nephron number and body weight or between nephron number and kidney size. From other studies, it appears that a greater than 2-fold variation is required in each of these parameters in order to demonstrate these relationships, suggesting they are highly dependent on scale. Our results are therefore not surprising since there was a less than 2-fold variation in each of the parameters examined. In summary, the relationship between nephron number and body or kidney size is most likely to be demonstrated when there is greater phenotypic variation either from genetic and/or environmental factors.     

Relationship between motoneuron number and myofibre number in the pig

A segment (at level T1) of spinal cord and a midbelly slice of the m. semitendinosus were removed from the largest- and smallest-birthweight littermate of each of 12 Large White pig litters sacrificed at various ages. When the ratio of myofibre numbers for large to small littermate within each litter was related to the ratio of motoneuron numbers then a significant correlation (r = 0.638, p less than 0.05) was observed. It is suggested that in utero growth retardation causes a reduction in myofibre numbers (within a litter), which in turn reduces motoneuron survival rate.     

Relationship between Fulton's condition factor and proximate body composition in three freshwater fish species

Morphometric‐based condition indices are widely used to assess proximate body composition and, collaterally, feeding and living conditions of fish. However, the exact relationship between condition indices and proximate body composition of fish and its relatedness to life history traits and seasonality has yet to be fully explored. Therefore, the aim of this study was to examine how the Fulton's condition factor (K‐factor) is related to the chemical composition (i.e. lipid, protein, water and carbon content, and molar carbon : nitrogen ratio), length and gonadal development of fish, and how these relationships are influenced by gender and seasonality in three freshwater fish species: Amur sleeper (Perccottus glenii), pumpkinseed (Lepomis gibbosus) and rudd (Scardinius erythrophthalmus). It was found that the strength and direction of association between the K‐factor and proximate body composition can vary markedly among fish species. The K‐factor correlated positively with gonadal development in pumpkinseed and Amur sleeper, while no such relationship existed in rudd. Condition factor can be a reliable measure of lipid content; however, the relationship was stronger in species with higher and more variable lipid contents. Moreover, a striking and consistently negative linkage was found between the K‐factor and water content of the fish body, which corresponds to the findings of several other studies. In turn, any relationship between the K‐factor and the protein content of fish was not detected. Gender seemed to exert a negligible effect on the relationship between the K‐factor and proximate body composition, while seasonal variance was obvious in most relationships.     

Variation in ovariole number and egg size of species of Dacus (Diptera; Tephritidae) and their relation to host specialization

Abstract. 1. An analysis of the number of ovarioles/ovary is presented for fourteen species of Dacus ranging from broad generalists (D.tryoni (Frogg.)) to strict host specialists (D.aglaia Hardy). For eight species data on egg size and the incubation time of eggs is also presented. Interrelationships among these life history variables and their relationships to host specialization were examined. 2. There was a broad positive relationship between the breadth of the natural host range (i.e. excluding cultivated fruits) and ovariole number (and hence potential fecundity). Highly polyphagous species had thirty-five to forty ovarioles/ovary while the specialists ranged from eight to twenty. 3. D.musae (Tryon), a specialist on banana, was an exception having an ovariole number similar to that of the most polyphagous species. 4. There was no clear relationship between egg size and ovariole number across eight species, though monophages were not fully represented in this analysis. Among these eight species there was no significant relationship between wing length and egg length, nor was there any such relationship among individuals within species. 5. There was considerable variation among species in the incubation time (at 25°C) for eggs, ranging from 24 h for D.cucumis French to 53 h for D.cacuminatus (Hering). This variation was only weakly related to egg size, though D.cucumis produced the largest eggs of any species examined. 6. Possible explanations for a relationship between potential fecundity and host range are discussed in terms of the balance between the abundance of host resources for generalists and their predictability for specialists. However, the observed relationship between ovariole number and host range must remain tentative, given that only fourteen species from one genus have been examined. Considerably more data is required for other dacines, for species in other tephritid subfamilies, and for other phytophagous insect groups to determine whether the relationship is generally applicable.