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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 derive 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.

Adaptation, Physiological↗

Nutritional requirements and diet choices of the pygmy rabbit (Brachylagus idahoensis): a sagebrush specialist.

Sagebrush (Artemisia tridentata) comprises up to 99% of the winter and 50% of the summer diets of pygmy rabbits (Brachylagus idahoensis). Few animals specialize on such plants as sagebrush, which contain high levels of plant chemicals that can be toxic. We investigated the nutritional requirements of pygmy rabbits and their ability and propensity to consume sagebrush alone and as part of a mixed diet. We compared diet choices of pygmy rabbits with that of a generalist forager, the eastern cottontail (Sylvilagus floridanus). Pygmy rabbits had a moderately low nitrogen requirement (306.5 mg N/kg(0.75)/d), but a relatively high energy requirement, needing 750.8 kJ digestible energy/kg(0.75)/d to maintain their body mass while residing in small cages. They digested fiber in pelleted diets similarly to other small hindgut fermenters, but both cottontails and pygmy rabbits digested the fiber in sagebrush better than expected based on its indigestible acid detergent lignin content. Pygmy rabbits consumed more sagebrush than cottontails, regardless of the amount and nutritional quality of supplemental pellets provided. When consuming sagebrush alone, they ate barely enough to meet their energy requirements, whereas cottontails ate only enough sagebrush to meet 67% of theirs. Both rabbit species ate more sagebrush as the quality and quantity of supplemental pellets declined, and more greenhouse-grown sagebrush than sagebrush grown outside. Urine was more acidic when consuming sagebrush than when consuming pellets, indicating detoxification by the liver. Pygmy rabbits do not require sagebrush to survive, but seem to tolerate terpenes and other plant chemicals in sagebrush better than cottontails do.

Animals↗

Serum LP(a) levels in African aboriginal Pygmies and Bantus, compared with Caucasian and Asian population samples.

Serum lipoprotein(a) (Lp(a)) and its correlates were studied in African Aboriginal Pygmies (n = 146) and Bantus (n = 208) from Cameroon. Geometric mean Lp(a) levels were 274 and 289 mg/l in Bantu males and females, respectively, and 220 and 299 mg/l in Pygmy males and females, the gender difference being significant in Pygmies (p = 0.024). In Pygmies 41% and 52% of the males and females, respectively, had Lp(a) levels above 300 mg/l, compared with 47% and 55% in Bantus. Overall, Lp(a) levels did not significantly differ between Pygmies and Bantus, and did not correlate with age, body mass index (BMI), systolic and diastolic blood pressure. Compared with healthy Asian and Caucasian population samples, age- and BMI-adjusted geometric Lp(a) means were 2.3- to 5.0-fold higher in Pygmy and Bantu males, and 2.9- to 3.6-fold higher in Pygmy and Bantu females (p < or = 0.05). Across the population samples studied ethnicity predicted 12% and 17% of serum Lp(a) variance in males and females, respectively.

Adult↗

Mother-infant interactions in slow lorises (Nycticebus bengalensis) and pygmy lorises (Nycticebus pygmaeus).

The study had three purposes: (1) to obtain information about mother-infant interactions in a rarely studied nocturnal prosimian, the pygmy loris (Nycticebus pygmaeus); (2) to compare pygmy lorises with a closely related and better-studied nocturnal prosimian, the Bengal slow loris (Nycticebus bengalensis); and (3) to determine how the presence of a second offspring affected mother-infant interactions in pygmy lorises. Three Bengal slow loris mothers and 3 pygmy loris mothers served as subjects, along with their 10 offspring (4 Bengal slow loris singletons, 2 pygmy loris singletons and 2 sets of pygmy loris twins). Observations were carried out in a zoo research facility for the first 24 weeks of the infants' lives. Although the two species differ in size and reproductive patterns, mother-infant interactions were similar. The primary modes of infant and adult contact were ventral and passive contact, respectively. Mothers parked their infants from the first week, and infants followed from the second week. Mothers displayed little protection or rejection, and there was little aggression. Infants solicited play and social grooming from their mothers. Pygmy loris mothers engaged in social grooming and play with their infants more frequently and for longer periods if the infant was a singleton rather than a twin.

Animals↗

Insulin-like growth factor I resistance in immortalized T cell lines from African Efe Pygmies.

Previous investigations suggested that resistance to GH was the cause of short stature of African Pygmies. Because many of the actions of GH are mediated by insulin-like growth factor I (IGF-I), we sought to determine whether Pygmy tissue was responsive to IGF-I. An initial effort to obtain HTLV-II-transformed T lymphoblast cell lines resulted in a single cell line that showed complete resistance to both IGF-I and GH in a clonal proliferation assay as well as decreased IGF-I binding. In the current study, we examined T cell lines from seven Efe Pygmy subjects, three neighboring Lese farmers, and six American controls and quantified clonal responses to IGF-I, GH, and insulin. The T cell lines from the Efe Pygmies were all completely resistant to the growth-promoting actions of IGF-I concentrations less than 250 micrograms/L and GH concentrations less than 500 micrograms/L. The Lese population, with whom there is admixture with the Efe population, showed heights and clonal responses to IGF-I and GH intermediate between those of Pygmies and American controls. The Pygmy T cell lines showed reduced clonal proliferation in response to high insulin concentrations known to act through the IGF-I receptor. These findings indicate that genetic IGF-I resistance is present in the T cell lines of Efe Pygmies and suggest that unresponsiveness to IGF-I may be responsible for their short stature.

Adult↗

Dissociation of systemic GH-IGF-I axis from a genetic basis for short stature in African Pygmies.

OBJECTIVES: To test the hypothesis that the primary cause of short stature in African Pygmies resides in low levels of insulin-like growth factor I (IGF-I), and to evaluate whether any observed alterations in their systemic IGF-I status can be dissociated from the effects of malnutrition and/or altered immune status. SETTING: Expedition to camps of partially sedentarized African Pygmies and neighbouring Bantu farmers living in the equatorial rain forest of Eastern Cameroon. DESIGN: Serum assays for hormonal and immune status were conducted for individuals (children, adolescents, young and old adults) showing no overt clinical nor biochemical signs of malnutrition, as judged from anthropometry, clinical examinations and serum assays of albumin, pre-albumin, retinol-binding protein, transferrin, iron, zinc, magnesium and calcium. RESULTS: African Pygmies did not differ from Europeans or Bantus in mean serum IGF-I concentrations, nor in the relationship between serum IGF-I and its major binding protein (IGFBP-3). However, although in both African groups the immunoglobulins IgG, IgM and IgE as well as the C-reactive protein and ceruloplasmin were above the normal range of European references, the Pygmies differed from their Bantu neighbours in their much higher IgG and IgM serum levels. A plot of serum IGF-I against these immunoglobulins in Pygmies revealed a reverse sigmoidal relationship, with the low IGF-I values associated with serum levels of IgG and IgM which clearly exceeded those found in the Bantus. CONCLUSIONS: This study indicates that in growing and adult African Pygmies showing no clinical nor biochemical signs of nutritional deficiency, serum IGF-I and IGFBP-3 (hence IGF-I bioavailability to its receptors) are essentially normal, and that low circulating levels of IGF-I in Pygmies reside in differential exposure and/or responsiveness to environmental challenge (e.g. infections) rather than in an inherited defect in the systemic growth-hormone (GH)-IGF-I axis.

Adolescent↗

Variation at 10 protein coding loci in the Mbenzele Pygmies from the Central African Republic and a comparison with microsatellite data.

Ten protein coding loci (6-PGD, A1-AT, ACP1, CaII, ESD, GC, GPX1, Hb beta, PGM1, and TF) were analyzed in the Mbenzele Pygmies from the Central African Republic. The frequency data were used to calculate the genetic distances between Mbenzele Pygmies and other African groups. In the principal coordinate plot of FST genetic distances, the Mbenzele cluster together with other Pygmies of the western cluster, the Biaka from C.A.R., Gielli from Cameroon, and Babinga from Congo. By contrast, they are considerably distanced from other Pygmy groups of the eastern cluster (Twa from Rwanda, Mbuti from Zaire). Genetic distances obtained using protein loci were compared with those based on microsatellite loci. The two distance matrices are insignificantly correlated (r = 0.268; one tail probability = 0.332), and the main difference is in the higher genetic affinity between the Mbenzele and Biaka Pygmies observed at the protein level. Although reasons underlying the discrepancy between inter-populational variation at protein and DNA loci are not established with certainty, the comparison suggests that the genetic distance between the Mbenzele and Biaka Pygmies at microsatellite loci could have been shaped by genetic drift.

Black People↗

Insulin-like growth factor-I unresponsiveness in an Efe Pygmy.

The cause of short stature in African Pygmies is unknown, but some evidence suggests that they are GH resistant. Since IGF-I mediates many actions of GH, we sought to determine if Pygmy tissue is responsive to IGF-I. We established HTLV-II-transformed cell lines from 1 Efe Pygmy, 1 African control, and 3 American controls, and quantified in vitro colony formation in response to IGF-I, GH, and insulin, and assessed IGF-I receptor binding. The Pygmy T-cell line showed no clonal responsiveness following stimulation with physiologic concentrations of IGF-I or any concentration of GH, but responded normally to insulin. IGF-I binding studies showed no binding to the Pygmy T-cell line with normal binding to control cells. The primary abnormality in this Pygmy T-cell line is IGF-I resistance at the receptor level with secondary GH resistance.

Adult↗

HLA class I variation in the West African Pygmies and their genetic relationship with other African populations.

We have studied the polymorphism of HLA class I in two West African Pygmy populations, namely, the Bakola from Cameroon and the Mbenzele from the Central African Republic. A unique number of HLA alleles and haplotypes showed specific patterns of these populations. In this study, we identify two alleles (B*37, B*41) and three haplotypes (A*30-B*37, A*66-B*41 and A*68-B*58) that appear to be 'private' or typical of Western Pygmies. These data reflect similarities with the AKA Pygmies from the Central African Republic. On the other hand, we failed to identify alleles that are found at high frequencies among other sub-Saharan populations (B*42, B*51). Allelic and haplotypic frequency distributions show differences between the two Pygmy groups, e.g. B*35 was very common in the Mbenzele but has been found to be absent in the Bakola. In contrast, B*53, which is found in the Bakola, has been found to be rare in the Mbenzele Pygmies. In order to analyse the genetic relationships of the Bakola and Mbenzele Pygmies with other sub-Saharan populations, HLA gene frequencies were subjected to the Neighbour-Joining tree analysis. The Mbenzele, Bakola and AKA were found to be relatively close to each other and isolated from other sub-African populations. However, both the genetic distances and the within-group variation suggests that the Bakola are more admixed with Bantu farmers than Mbenzele.

Africa, Western↗

Pygmy mouse fibroblasts in tissue culture: evaluation of multiplication stimulating activity (MSA) receptors and growth responses to MSA.

The pygmy mouse has been proposed as a model for growth hormone resistance; it has normal serum somatomedin levels and does not respond to growth hormone treatment. In order to determine if the growth impairment is caused by a defect in somatomedin binding or in postreceptor action of somatomedin we compared fibroblasts derived from pygmy mice with those from normal appearing littermates. Using multiplication-stimulating activity (MSA) as a model somatomedin we found a normal Ka of binding to the cell surface MSA receptor but a significantly increased number of MSA receptors on the fibroblasts derived from pygmy mice. Studies of thymidine incorporation into DNA failed to demonstrate a difference between pygmy and normal fibroblasts in their responses to MSA alone, but there was a significantly greater thymidine incorporation into the DNA of normal fibroblasts when both competence factor (platelet-derived growth factor) and progression factors (somatomedins and growth hormone deficient platelet-poor plasma) were present in the test medium. On the other hand, cell proliferation studies did not demonstrate a consistent difference in the growth rate of normal versus pygmy fibroblasts. The data support the conclusion that the imparied growth of the pygmy mouse in vivo may be caused by factors which lie outside of the growth hormone-somatomedin pathway.

Animals↗

Molecular epidemiology of 58 new African human T-cell leukemia virus type 1 (HTLV-1) strains: identification of a new and distinct HTLV-1 molecular subtype in Central Africa and in Pygmies.

To gain new insights on the origin, evolution, and modes of dissemination of human T-cell leukemia virus type I (HTLV-1), we performed a molecular analysis of 58 new African HTLV-1 strains (18 from West Africa, 36 from Central Africa, and 4 from South Africa) originating from 13 countries. Of particular interest were eight strains from Pygmies of remote areas of Cameroon and the Central African Republic (CAR), considered to be the oldest inhabitants of these regions. Eight long-term activated T-cell lines producing HTLV-1 gag and env antigens were established from peripheral blood mononuclear cell cultures of HTLV-1 seropositive individuals, including three from Pygmies. A fragment of the env gene encompassing most of the gp21 transmembrane region was sequenced for the 58 new strains, while the complete long terminal repeat (LTR) region was sequenced for 9 strains, including 4 from Pygmies. Comparative sequence analyses and phylogenetic studies performed on both the env and LTR regions by the neighbor-joining and DNA parsimony methods demonstrated that all 22 strains from West and South Africa belong to the widespread cosmopolitan subtype (also called HTLV-1 subtype A). Within or alongside the previously described Zairian cluster (HTLV-1 subtype B), we discovered a number of new HTLV-1 variants forming different subgroups corresponding mainly to the geographical origins of the infected persons, Cameroon, Gabon, and Zaire. Six of the eight Pygmy strains clustered together within this Central African subtype, suggesting a common origin. Furthermore, three new strains (two originating from Pygmies from Cameroon and the CAR, respectively, and one from a Gabonese individual) were particularly divergent and formed a distinct new phylogenetic cluster, characterized by specific mutations and occupying in most analyses a unique phylogenetic position between the large Central African genotype (HTLV-1 subtype B) and the Melanesian subtype (HTLV-1 subtype C). We have tentatively named this new HTLV-1 genotype HTLV-1 subtype D. While the HTLV-1 subtype D strains were not closely related to any known African strain of simian T-cell leukemia virus type 1 (STLV-1), other Pygmy strains and some of the new Cameroonian and Gabonese HTLV-1 strains were very similar (>98% nucleotide identity) to chimpanzee STLV-1 strains, reinforcing the hypothesis of interspecies transmission between humans and monkeys in Central Africa.

Africa, Central↗

Population genetics of red cell enzymes in Pygmies: a conclusive account.

In the course of a long-term research project, three groups of Pygmies and some non-Pygmy Central Africans have been examined for the following red cell enzyme markers: ACP, PGM1, PGM2, PEPA, PEPB, and PEPC, AK, ADA, and PHI. Several other red cell enzymes (ESD, CA1 and CA2, GPT, GLO, and DIA1) have been studied in only some of these groups. This paper reports all the information we obtained, including what we have already published. The following conclusions can be drawn from the whole body of data: (1) Gene patterns of Pygmies are those typical of other Africans (e.g.: lack of ADA2 and AK2 genes, low GPT2 gene frequency, polymorphism of the CA2 locus, and presence at polymorphic frequencies of PEPA2 allele. (2) Superimposed on this African genetic makeup, a number of Pygmy characters were identified, namely, a private polymorphism for the PGM26 Pygmy allele and possibly one for the PEPC2 allele, and particularly high ACPR and low PGM12 gene frequencies. (3) Some markers, especially PGM1 and ACP, turned out also to discriminate efficiently among different groups of Pygmies.

Black People↗

Binary and microsatellite polymorphisms of the Y-chromosome in the Mbenzele pygmies from the Central African Republic.

This study analyzes the variation of six binary polymorphisms and six microsatellites in the Mbenzele Pygmies from the Central African Republic. Five different haplogroups (B2b, E(xE3a), E3a, P and BR(xB2b,DE,P)) were observed, with frequencies ranging from 0.022 (haplogroup P) to 0.609 (haplogroup E3a). A comparison of haplogroup frequencies indicates a close genetic affinity between the Mbenzele and the Biaka Pygmies, a finding consistent with the common origin and the geographical proximity of the two populations. The haplogroups P, BR(xB2b,DE,P) and E(xE3a), which are rare in sub-Saharan Africa but common in western Eurasia, were observed with frequencies ranging from 0.022 (haplogroup P) to 0.087 (haplogroup E(xE3a)). Thirty different microsatellite haplotypes were detected, with frequencies ranging from 0.022 to 0.152. The Mbenzele share the highest percent of microsatellite haplotypes with the Biaka Pygmies. Five out seven haplotypes which are shared by the Mbenzele and Biaka Pygmies belong to haplogroup E3a, which suggests that they are of Bantu origin. The plot based on F(st) genetic distances calculated using microsatellite data provides a picture of population relationships which is in part congruent and in part complementary to that obtained using haplogroup frequencies. Finally, the Mbenzele and Biaka Pygmies were found to be markedly more genetically similar using Y-chromosomal than autosomal microsatellites. We suggest that this could be due to the higher phylogenetic stability of Y-chromosome and to the effect of the male-biased gene flow during the Bantu expansion.

Alleles↗

Polymorphism and mapping of the IGF1 gene, and absence of association with stature among African Pygmies.

Probes detecting restriction fragment length polymorphisms (RFLPs) in the insulinlike growth factor (IGF1) gene were isolated and allele frequencies in different human populations determined. No difference was detected between the distribution of IGF1 alleles in Pygmies versus non-Pygmy black Africans, despite the proposal that a defect in the IGF1 gene might be responsible for Pygmy short stature. This was supported by the absence of a correlation of IGF1 genotype with height in the C.A.R. Pygmies. Polymerase chain reaction (PCR) and direct sequencing failed to demonstrate an alteration in the region upstream the IGF1 start site in Pygmies. Linkage analysis demonstrated that IGF1 is tightly linked to the phenylalanine hydroxylase gene on chromosome 12q22-24.1.

Africa↗

Spontaneous symbol acquisition and communicative use by pygmy chimpanzees (Pan paniscus).

Two pygmy chimpanzees (Pan paniscus) have spontaneously begun to use symbols to communicate with people. In contrast to common chimpanzees (Pan troglodytes) using the same communicative system, the pygmy chimpanzees did not need explicit training in order to form referential symbol-object associations. Instead, they acquired symbols by observing others use these symbols in daily communications with them. In addition, the pygmy chimpanzees have begun to comprehend spoken English words and can readily identify lexigrams upon hearing the spoken words. By contrast, common chimpanzees who received similar exposure to spoken English are unable to do so. The older pygmy chimpanzee has begun to form requests of the form agent-verb-recipient in which he is neither the agent nor the recipient. By contrast, similarly aged common chimpanzees limited their requests to simple verbs, in which the agent was always presumed to be the addressee and the chimpanzee itself was always the recipient, thus they had no need to indicate a specific agent or recipient. These results suggest that these pygmy chimpanzees exhibit symbolic and auditory perceptual skills that are distinctly different from those of common chimpanzees.

Animal Communication↗

Dwarfism in the pygmy. An isolated deficiency of insulin-like growth factor I.

Insulin-like growth factors (IGFs) I and II, which are present in normal human beings, were measured in serum samples from 11 pygmies from the Central African Republic, 31 controls, and 12 patients with growth hormone deficiency. The mean serum concentration of IGF-I (+/- S.E.M.) was 68.6 +/- 8 ng per milliliter in pygmies, as compared with 193 +/- 10 ng per milliliter in controls (P less than 0.001) and 24 +/- 4 ng per milliliter in patients with growth hormone deficiency (P less than 0.05). Mean serum concentrations of IGF-II in controls, pygmies, and growth hormone-deficient patients were 647 +/- 22, 503 +/- 37, and 252 +/- 29 ng per milliliter, respectively. The serum IGF-I concentration was within the normal range in only one pygmy, whereas IGF-II values were within the normal range in 10 of 11. Pygmies appear to have a major defect in the production of IGF-I.

Adolescent↗

Short KIR haplotypes in pygmy chimpanzee (Bonobo) resemble the conserved framework of diverse human KIR haplotypes.

Some pygmy chimpanzees (also called Bonobos) give much simpler patterns of hybridization on Southern blotting with killer cell immunoglobulin-like receptor (KIR) cDNA probes than do either humans or common chimpanzees. Characterization of KIRs from pygmy chimpanzees having simple and complex banding patterns identified nine different KIRs, representing seven genes. Five of these genes have orthologs in the common chimpanzee, and three of them (KIRCI, KIR2DL4, and KIR2DL5) also have human orthologs. The remaining two genes are KIR3D paralogous to the human and common chimpanzee major histocompatibility complex A- and/or -B-specific KIRs. Within a pygmy chimpanzee family, KIR haplotypes were defined. Simple patterns on Southern blot were due to inheritance of "short" KIR haplotypes containing only three KIR genes, KIRCI, KIR2DL4, and KIR3D, each of which represents one of the three major KIR lineages. These three genes in pygmy chimpanzees or their corresponding genes in humans and common chimpanzees form the conserved "framework" common to all KIR haplotypes in these species and upon which haplotypic diversity is built. The fecundity and health of individual pygmy chimpanzees who are homozygotes for short KIR haplotypes attest to the viability of short KIR haplotypes, indicating that they can provide minimal, essential KIRs for the natural killer and T cells of the hominoid immune system.

Amino Acid Sequence↗

HTLV-II seroprevalence in pygmies across Africa since 1970.

HTLV-II-specific antibodies, with patterns similar to those in the Americas, were present in sera collected about 1970 from Bambuti pygmies in Zaire (14/102; 14%) and from pygmies in Cameroon (5/214; 2.3%), and were more prevalent than HTLV-I. In the Central African Republic, 504 pygmies were HTLV negative. After finding of 4 HTLV-II seropositives among 12 Bambuti pygmies sampled in 1991, this established that HTLV-II or a related retrovirus is present as an ancient endemic in some, but not all, insulated groups of African pygmies, similar to the HTLV-II distribution in Amerindian populations. The endemic among the oldest inhabitants of central Africa, and the occasional and scattered occurrence of apparent HTLV-II among predominant HTLV-I in other Africans, fit well with an ancient African virus and not with importation from the New World. Theories on the origin and evolution of the primate T-lymphotropic viruses (PTLVs) should take into account the longstanding presence of HTLV-II-type viruses in both the Old and New World. Present serology suggests identity of the African viruses with HTLV-II, but their assignment to a new HTLV type is open should genetic analysis show strong divergence from American HTLV-II. Clinical expression, if any, remains to be studied.

Blotting, Western↗