Multiple independent appearances of the cecal appendix in mammalian evolution and an investigation of related ecological and anatomical factors

Although the cecal appendix has been widely viewed as a vestige with no known function or a remnant of a formerly utilized digestive organ, the evolutionary history of this anatomical structure is currently unresolved. A database was compiled for 361 mammalian species, and appendix characters were mapped onto a consensus phylogeny along with other gastrointestinal and behavioral characters. No correlation was found between appearance of an appendix and evolutionary changes in diet, fermentation strategy, coprophagia, social group size, activity pattern, cecal shape, or colonic separation mechanism. Appendix presence and size are positively correlated with cecum and colon size, even though this relationship rests largely on the larger size of cecum and colon in taxa that have an appendix. The appendix has evolved minimally 32 times, but was lost fewer than seven times, indicating that it either has a positive fitness value or is closely associated with another character that does. These results, together with immunological and medical evidence, refute some of Darwin's hypotheses and suggest that the appendix is adaptive but has not evolved as a response to any particular dietary or social factor evaluated here.     

Genetic constraints on phenotypic evolution in Nigella (Ranunculaceae)

The present investigation examines the role of genetic constraints in shaping evolutionary change in the Nigella arvensis species complex. Parent-offspring analyses of two populations of N. degenii demonstrated high heritabilities for a wide range of vegetative and floral characters, indicating a great potential for further adaptive change. The populations differed significantly in the heritability for leaf length and in the genetic correlation between plant height and peduncle length, suggesting that these populations would respond differently to identical selection pressures. There was a tendency for large-scale diversity to extrapolate within-population variability, at least for the floral trait associations, while genetic data from a segregating F3 hybrid population indicated stability in the correlation structure across two environments. On the basis of hybrid data, I propose that inbreeding effects and pleiotropic relationships with leaf size may have facilitated the reduction in floral morphology accompanying the evolution of autogamy in related taxa.     

Comparative anatomy and phylogenetic distribution of the mammalian cecal appendix

A recently improved understanding of gut immunity has merged with current thinking in biological and medical science, pointing to an apparent function of the mammalian cecal appendix as a safe‐house for symbiotic gut microbes, preserving the flora during times of gastrointestinal infection in societies without modern medicine. This function is potentially a selective force for the evolution and maintenance of the appendix, and provides an impetus for reassessment of the evolution of the appendix. A comparative anatomical approach reveals three apparent morphotypes of the cecal appendix, as well as appendix‐like structures in some species that lack a true cecal appendix. Cladistic analyses indicate that the appendix has evolved independently at least twice (at least once in diprotodont marsupials and at least once in Euarchontoglires), shows a highly significant (P < 0.0001) phylogenetic signal in its distribution, and has been maintained in mammalian evolution for 80 million years or longer.     

Observations on Variations in the Male Sex Characters and Their Relation to the Androgenic Gland in the Shrimp Macrobrachium idae (Heller)

A majority of the males of the brackish water shrimp, Macrobrachium idae, show a hypertrophy in the length of the second cheliped, the length and spinuosity of the appendix masculina and the width of the vas deferens, during the breeding season. Statistical analyses of the measurements of these characters show that this hypertrophy has only very little correlation to the size of the individual, and that the above sex characters have marked variation in a population having the same body length. The hypertrophy of these structures is more linked with the extent of development of the androgenic gland. Also, in such individuals, the androgenic gland shows signs of increased secretory activity as evidenced by increase in size of the cells, vacuolisation of cytoplasm and presence of degenerative areas. It is suggested that there is a direct relation between the secretory activity of the androgenic gland and the hypertrophy of these structures i.e. there is a “cause and effect” relation between the androgenic gland and the primary and secondary sex characters.     

TEMPORAL VARIATION OF FLOWER AND FRUIT SIZE IN RELATION TO SEED YIELD IN CELANDINE POPPY (CHELIDONIUM MAJUS; PAPAVERACEAE)

Ecological and evolutionary studies typically consider variation in single reproductive characters in isolation, without considering how they might be correlated with other reproductive and vegetative characters. In our study, we examined temporal patterns of variation and correlation in flower diameter and fruit length during a reproductive phase in two Massachusetts populations of the herb, Chelidonium majus. We also examined the relationships of such variation to measurements of seed yield components (mean seed weight and number per fruit) and aspects of plant vegetative size. Most of the variation in the sizes of reproductive characters occurred within individual plants, instead of among plants or between populations. Flower and fruit sizes as well as seed number per fruit declined significantly during the season in both populations. Only mean seed size per fruit was relatively stable for individual plants in both populations. Conserving resources by a gradual reduction in the size of reproductive characters over the season may be a strategy for maternal plants to continue seed production. The strong, persistent patterns of correlation between certain characters, such as flower and fruit size, in spite of extensive phenotypic plasticity, was interpreted as indirect evidence for developmental correlation. Furthermore, vegetatively larger plants produced not only more flowers and fruits, but also consistently larger flowers and fruits. The results emphasize that variation in fitness characters, such as seed size and number, should not be viewed in isolation from vegetative characters, flower, and fruit sizes in ecological and evolutionary studies, if the goal is to understand the mechanisms of natural selection in wild populations.     

Effect of Bacterial Contamination on Cecal Size and Cecal Contents of Gnotobiotic Rodents

In the present investigation the effect of various bacterial contaminations of gnotobiotic mice and rats on cecal size is presented. Of the species tested, Bacteroides oralis and Fusobacterium nucleatum did not establish in germ-free mice. Streptococcus mutans, Clostridium difficile, a Neisseria strain and two recent cecal isolates established, but failed to exert an effect upon the cecum of mice. A group K streptococcus and B. fragilis increased the cecal size apparently by increasing the levels of water-soluble protein, peptides, and carbohydrates in the cecal contents. Mixed ileal bacteria decreased the cecal size by preventing accumulation of soluble proteins and carbohydrates in the cecum. A Peptococcus strain caused a reduction by lowering the levels of insoluble material in the cecum. When this strain was combined with two Clostridium isolates and introduced into gnotobiotic rats, 50 to 65% cecal reduction was observed. This polycontamination did not decrease the per cent water of the cecal contents but caused lower levels of both soluble and insoluble material to accumulate in the cecum. No net nitrogen absorption from the distal small intestine occurred in either the germ-free or polycontaminated rats.     

Morphological variation in Puya (Bromeliaceae): an allometric study

A group of representative species of the genus Puya was studied to determine if there are allometric relationships between vegetative and floral parts, whether these relationships correlate with their pollination system and if plant size is correlated with elevation and latitude. Fifty-three species representing the morphological variation and distribution of the genus were studied. Total plant height, as well as leaf, inflorescence, petal and sepal length were measured and these data subjected to univariate and multivariate analyses. To test for correlation between plant size and altitudinal and latitudinal distribution, ANOVAs were performed. When the pollination system of a species is known, additional multivariate and univariate analyses were also performed. The results indicate that the characters studied are correlated with a size component, exhibiting positive allometry for sepal and petal length and negative allometry for leaf length. Inflorescence length is an isometric character. There was no significant correlation between plant size and altitudinal or latitudinal distribution. The ANOVAs show that the only character correlated to pollinator type was petal length. Small plants with small flowers are correlated to pollinators such as insects, while medium to large plants with medium to large-sized flowers are correlated with pollinators such as birds and bats. Large plants have small flowers, that are more evident and attract more pollinators.     

Range size in North American Enallagma damselflies correlates with wing size

1. Cross-species macroecological comparisons in freshwater invertebrates have been restricted by a lack of large-scale distributional data, and robust phylogenies. Here, we use data from the OdonataCentral database to explore body length–range size and wing length–range size relationships in damselflies from the genus Enallagma ; the recent publication of a phylogeny for this group meant that, as well as a cross-species analysis, we were able to assess relationships in a phylogenetically controlled manner.
2. For cross-species comparisons, only wing length showed significant (positive) regression relationships with range size and occupancy, although the inclusion of body length in multiple regressions increased the fit of the models. Damselflies with larger wings relative to their body length had larger distributions, a result confirmed by a significant positive relationship between range size and residuals from the regression of wing size on body size.
3. For the phylogenetically controlled analyses, only wing length contrast scores were significantly related to distribution patterns and entered into regression models; the significant positive relationships between wing length contrasts and both range size and occupancy contrasts suggested that evolutionary increases in wing length had occurred alongside range expansions.
4. Together these results suggest that species of Enallagma with larger wings (both absolute and relative to body length) tend to be more widely distributed in North America and that the evolution of wing size may have played a role in range expansion. No such relationships were evident for body size. We discuss the potential importance of wing morphometrics for studying the evolutionary ecology of freshwater insects.     

Enterotoxemia in rabbits.

The presence of Clostridium perfringens Type E iota toxin was confirmed from the cecal contents of 23 of 46 rabbits which died of enteritis complex. The most consistent lesions observed were hemorrhage and edema in the cecum. Rabbit toxicity tests showed the toxic cecal contents were lethal for young rabbits unless incubated with Clostridium perfringens Type E antiserum.     

Lowering sample size in comparative analyses can indicate a correlation where there is none: example from Rensch's rule in primates

The fact that characters may co-vary in organism groups because of shared ancestry and not always because of functional correlations was the initial rationale for developing phylogenetic comparative methods. Here we point out a case where similarity due to shared ancestry can produce an undesired effect when conducting an independent contrasts analysis. Under special circumstances, using a low sample size will produce results indicating an evolutionary correlation between characters where an analysis of the same pattern utilizing a larger sample size will show that this correlation does not exist. This is the opposite effect of increased sample size to that expected; normally an increased sample size increases the chance of finding a correlation. The situation where the problem occurs is when co-variation between the two continuous characters analysed is clumped in clades; e.g. when some phylogenetically conservative factors affect both characters simultaneously. In such a case, the correlation between the two characters becomes contingent on the number of clades sharing this conservative factor that are included in the analysis, in relation to the number of species contained within these clades. Removing species scattered evenly over the phylogeny will in this case remove the exact variation that diffuses the evolutionary correlation between the two characters - the variation contained within the clades sharing the conservative factor. We exemplify this problem by discussing a parallel in nature where the described problem may be of importance. This concerns the question of the presence or absence of Rensch's rule in primates.     

Variation in growth of overwintered stolons of contrasting white clover populations in response to temperature, photoperiod and spring environment

The response of overwintered stolons of nine contrasting white clover populations to temperature, photoperiod and natural conditions was studied in six environments during the spring. Rate of leaf appearance, leaflet length, petiole length, stolon internode length, dry matter distribution within the plant and total dry weight were measured on 15 plants of each population/environment combination. Most characters, except leaf size and proportion of dry matter allocated to leaf, responded positively to temperature in the range 10 – 20°C. A positive effect of photoperiod extension was also found for all characters except rate of leaf appearance, internode length and distribution of dry matter to leaf. Population differences in response to environment were found which were related to both leaf size classification and origin. Stolon dry weight was positively correlated with leaf length, petiole length and stolon internode length in most environments. The relationships between the eight characters were often complex and canonical variate analysis provided a convenient way to discriminate between the populations based on all eight characters.     

Histology of vermiform appendix-like organ in slow loris.

The vermiform appendix-like organ (VALO) of the slow loris was investigated for its histology and immunohistochemical characteristics. The VALO has a much thinner wall with flat folded mucosa and shallower crypts than the cecal mucosa, while cellular components and population of the mucosa were similar to those of the cecum. No coalescent lymph nodules were seen in the submucosa. Immunohistochemically 5-HT-positive cells in the crypts and CD3- and CD8-positive lymphocytes in the lymph nodules were shown in the VALO as well as in the cecum. These findings suggest that the VALO is a low-differentiated vermiform appendix of the slow loris.     

The integrated phenotype

Proper functioning of complex phenotypes requires that multiple traits work together. Examination of relationships among traits within and between complex characters and how they interact to function as a whole organism is critical to advancing our understanding of evolutionary developmental plasticity. Phenotypic integration refers to the relationships among multiple characters of a complex phenotype, and their relationships with other functional units (modules) in an organism. In this review, I summarize a brief history of the concept of phenotypic integration in plant and animal biology. Following an introduction of concepts, including modularity, I use an empirical case-study approach to highlight recent advance in clarifying the developmental and genomic basis of integration. I end by highlighting some novel approaches to genomic and epigenetic perturbations that offer promise in further addressing the role of phenotypic integration in evolutionary diversification. In the age of the phenotype, studies that examine the genomic and developmental changes in relationships of traits across environments will shape the next chapter in our quest for understanding the evolution of complex characters.     

Effects of two probiotic Lactobacillus strains on jejunal and cecal microbiota of broiler chicken under acute heat stress condition as revealed by molecular analysis of 16S rRNA genes

We examined the effects of probiotic Lactobacillus strains of Lactobacillus agilis JCM 1048 and Lactobacillus salivarius subsp. salicinius JCM 1230 on jejunal and cecal microbiota of broiler chicken under heat stress condition using terminal restriction fragment length polymorphism (T-RFLP) analysis. The jejunal bacterial community was limited to a few bacterial groups, mostly Lactobacillus spp. A relatively abundant and higher prevalence of Lactobacillus spp. were observed in the jejunal and cecal microbiota of the probiotic chickens compared with those of the control chickens under heat stress condition. In general, the probiotic strains did not significantly affect the abundance of L. agilis and L. salivarius in chicken intestine but clearly contributed to increasing their prevalence in the probiotic chickens. The probiotic Lactobacillus strains enriched the diversity of Lactobacillus flora in chicken jejunum and cecum by increasing the abundance and prevalence of Lactobacillus spp. inhabiting the intestine. The richness of Lactobacillus species tended to be similar among the jejunal and cecal microbiota. The bacterial community of cecum was complex and age-dependent. The major components of the cecal microbiota were clostridia and lactobacilli. The Clostridium subcluster XIVa was the most predominant group in chicken cecum. Probiotic Lactobacillus strains restored the microbial balance and maintained the natural stability of indigenous bacterial microbiota following heat stress-induced changes.     

Distinct evolutionary patterns of morphometric sperm traits in passerine birds

The striking diversity of sperm shape across the animal kingdom is still poorly understood. Postcopulatory sexual selection is an important factor driving the evolution of sperm size and shape. Interestingly, morphometric sperm traits, such as the length of the head, midpiece and flagellum, exhibit a strong positive phenotypic correlation across species. Here we used recently developed comparative methods to investigate how such phenotypic correlations between morphometric sperm traits may evolve. We compare allometric relationships and evolutionary trajectories of three morphometric sperm traits (length of head, midpiece and flagellum) in passerine birds. We show that these traits exhibit strong phenotypic correlations but that allometry varies across families. In addition, the evolutionary trajectories of the midpiece and flagellum are similar while the trajectory for head length differs. We discuss our findings in the light of three scenarios accounting for correlated trait evolution: (i) genetic correlation; (ii) concerted response to selection acting simultaneously on different traits; and (iii) phenotypic correlation between traits driven by mechanistic constraints owing to selection on sperm performance. Our results suggest that concerted response to selection is the most likely explanation for the phenotypic correlation between morphometric sperm traits.     

On the relationship between body dimensions and appendix length

In the last decades a lot of significant correlations between clinical and anthropometric parameters were found, e.g. the relationships between somatotypical and endocrine parameters or between body dimensions and echocardiographic heart size. During operation, surgeons often find a wide range of appendix size of patients with varying body dimensions. A goal of this work is the detection of the relationships between anthropometric characteristics (height, weight, body mass index (= BMI)), inflammation parameters (histological inflammation degree, leukocyte count and C-reactive protein (= CRP)) and the appendix length with n=167 consecutive patients (time period 1.1.2004 - 31.12.2006; range of age 7-95 years, median age 20 years; average age of men 27.7, women 26.6 years), which had to be operated because of acute appendicitis. Appendectomy was laparoscopic in 95 % of the patients and open in 5 %. The appendix length of the male patients was high-significantly longer with a mean of 7.5 cm (SD = 2.1) than those of the female patients with 6.3 cm (SD = 1.6). The following mean anthropometric values were registered: height 169.8 (SD = 15.6) cm for the males and 165.2 (SD = 9.6) cm for the females, weight 70.4 (SD = 23.4) kg for the males and 61.9 (SD = 16.1) kg for the females, BMI 24.1 (SD = 5.4) kg/m2 for the males and 22.9 (SD = 4.8) kg/m2 for the females. The mean leukocyte count was 12,700 (SD = 5500)/ml for the males and 11,600 (SD = 4900)/ml for the females, CRP was 4.3 (SD = 5.4) mg/l for the males and 4.5 (SD = 8.9) mg/l for the females. Concerning the histopathological diagnosis, n=76 patients (45.5 %) suffered from a submucous fibrosis bliteration, n 1 (12.6 ) from an acute purulent inflammation degree, n = 65 (38.9 ) from an ulcerative hlegmonous inflammation degree and n (3 ) from a perforation. While no significant Spearman's correlation coefficient between appendix length and CRP was found, the appendix length correlated highly significantly to the body weight (r = 0.25) and significantly to leukocyte count (r = 0.16), body height (r = 0.16), BMI (r = 0.17) and the histological inflammation degree (r = 0.18). Thus, the correlations found were lower than the relationships between anthropometric and endocrine or echocardiographic parameters described in literature.     

Quantitative detection of Clostridium perfringens in the broiler fowl gastrointestinal tract by real-time PCR

Strains of Clostridium perfringens are a frequent cause of food-borne disease and gas gangrene and are also associated with necrotic enteritis in chickens. To detect and quantify the levels of C. perfringens in the chicken gastrointestinal tract, a quantitative real-time PCR assay utilizing a fluorogenic, hydrolysis-type probe was developed and utilized to assay material retrieved from the broiler chicken cecum and ileum. Primers and probe were selected following an alignment of 16S rDNA sequences from members of cluster I of the genus Clostridium, and proved to be specific for C. perfringens. The assay could detect approximately 50 fg of C. perfringens genomic DNA and approximately 20 cells in pure culture. Measurements of the analytical sensitivity determined with spiked intestinal contents indicated that the consistent limit of detection with ileal samples was approximately 10(2) CFU/g of ileal material, but only about 10(4) CFU/g of cecal samples. The decreased sensitivity with the cecal samples was due to the presence of an unidentified chemical PCR inhibitor(s) in the cecal DNA purifications. The assay was utilized to rapidly detect and quantify C. perfringens levels in the gut tract of broiler chickens reared without supplementary growth-promoting antibiotics that manifested symptoms of necrotic enteritis. The results illustrated that quantitative real-time PCR correlates well with quantification via standard plate counts in samples taken from the ileal region of the gastrointestinal tract.     

家兔盲肠和阑尾淋巴管的研究

用色素穿刺注入法、动脉内墨汁硝酸银水溶液注入法研究了家兔盲肠和阑尾淋巴管的起始、走向及所属淋巴结。盲肠和阑尾的毛细淋巴营均起始于固有膜,后依次穿过肠壁各层移行为系膜内淋巴管。后者经阑尾、盲肠淋巴结与肠系膜上淋巴结群中的相应淋巴结相联系。盲肠粘膜下层淋巴管内开始出现瓣膜;阑尾淋巴管瓣膜则在阑尾系膜缘内才出现。     

Plasticity or fixed adaptive traits? Strategies for predation avoidance in Rana arvalis tadpoles

Tadpoles of Rana arvalis originating from seven island populations were tested for responses to non-lethal predator presence. In general, tadpole growth was reduced and the relative tail depth was increased at predator presence. There was no effect of predator presence on the predicted size at metamorphosis. The differentiation rate, translating as length of the larval period, was lower at predator presence, but this seems to be merely an effect of the reduced growth. Although populations differed with respect to growth, relative tail length, relative tail depth, differentiation rate and predicted size at metamorphosis, no obvious differences were found in their responses to predator presence. Data on predator occurrences in the source ponds show that tadpoles originating from ponds with a high predation pressure have a higher differentiation rate, i.e. they will metamorphose at an earlier date than those from “safe” ponds (if raised under the same conditions). Moreover, they are also predicted to metamorphose at a smaller size, which is in accordance with theoretical models. Despite the fact that populations differed in growth, no correlation was found between growth and predation risk in the source ponds. Received: 16 March 1998 / Accepted: 18 July 1998