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Grandmothers and the evolution of human longevity.

Great apes, our closest living relatives, live longer and mature later than most other mammals and modern humans are even later-maturing and potentially longer-lived. Evolutionary life-history theory seeks to explain cross-species differences in these variables and the covariation between them. That provides the foundation for a hypothesis that a novel role for grandmothers underlies the shift from an ape-like ancestral pattern to one more like our own in the first widely successful members of genus Homo. This hypothesis links four distinctive features of human life histories: 1). our potential longevity, 2). our late maturity, 3). our midlife menopause, and 4). our early weaning with next offspring produced before the previous infant can feed itself. I discuss the problem, then, using modern humans and chimpanzees to represent, respectively, genus Homo and australopithecines, I focus on two corollaries of this grandmother hypothesis: 1). that ancestral age-specific fertility declines persisted in our genus, while 2). senescence in other aspects of physiological performance slowed down. The data are scanty but they illustrate similarities in age-specific fertility decline and differences in somatic durability that are consistent with the hypothesis that increased longevity in our genus is a legacy of the "reproductive" role of ancestral grandmothers.

Aging↗

Functions of corticotropin-releasing hormone in anthropoid primates: from brain to placenta.

Corticotropin-releasing hormone (CRH) is an ancient regulatory molecule. The CRH hormone family has at least four ligands, two receptors, and a binding protein. Its well-known role in the hypothalamic-pituitary-adrenal (HPA) axis is only one of many. The expression of CRH and its related peptides is widespread in peripheral tissue, with important functions in the immune system, energy metabolism, and female reproduction. For example, CRH is involved in the implantation of fertilized ova and in maternal tolerance to the fetus. An apparently unique adaptation has evolved in anthropoid primates: placental expression of CRH. Placental CRH stimulates the fetal adrenal zone, an adrenal structure unique to primates, to produce dehydroepiandrosterone sulfate (DHEAS), which is converted to estrogen by the placenta. Cortisol induced from the fetal and maternal adrenal glands by placental CRH induces further placental CRH expression, forming a positive feedback system that results in increasing placental production of estrogen. In humans, abnormally high placental expression of CRH is associated with pregnancy complications (e.g., preterm labor, intrauterine growth restriction (IUGR), and preeclampsia). Within anthropoid primates, there are at least two patterns of placental CRH expression over gestation: monkeys differ from great apes (and humans) by having a midgestational peak in CRH expression. The functional significance of these differences between monkeys and apes is not yet understood, but it supports the hypothesis that placental CRH performs multiple roles during gestation. A clearer understanding of the diversity of patterns of placental CRH expression among anthropoid primates would aid our understanding of its role in human pregnancy.

Animals↗

Episodic memory in primates.

Episodic memory refers to a system of memory with the capacity to recollect specific events from an individual's life. Some psychologists have suggested that episodic memory is a uniquely human phenomenon. We challenge that idea and present evidence that great apes and other primates may possess episodic-like memory. We review criteria developed to assess episodic-like memory in nonhumans, and how they apply to primates. In particular, we discuss the criteria of Clayton et al. [2001], who stated that episodic-like memory is based on the retrieval of multiple and integrated components of an event. We then review eight studies examining memory in great apes and apply the Clayton et al. criteria to each of them. We summarize the evidence that is compatible with the existence of episodic-like memory, although none of the data completely satisfy the Clayton et al. criteria. Morover, feelings of pastness and feelings of confidence, which mark episodic memory in humans, have not been empirically addressed in nonhuman primates. Future studies should be directed at these aspects of memory in primates. We speculate on the functional significance of episodic memory in nonhuman primates.

Animals↗

Human cognitive ecology: an instructive framework for comparative primatology.

In this review, research on human cognitive ecology is compared with studies of the cognitive ecologies of apes-especially the common chimpanzee. The objective was to assess the feasibility of extending an activity-theory framework developed in studies of humans to an integrated approach for studying the cognitive accomplishments and skills of other primates living in the wild. Six generalizations were abstracted from studies of humans: 1) Social and material environments are arranged to facilitate production. 2) Human activity is shaped by conceptual and cultural principles that provide underlying logic for working knowledge and practice. 3) Schemata (multimodal, mental representations of procedures, strategies, and techniques) govern performance in a domain. 4) Working knowledge, skills, and social identities are co-constructed in communities of practice. 5) Rehearsal improves skilled performances, from which reputations as well as material products are derived. 6) Planning and emergence are in productive tension in human practices. These generalizations are applied to findings in the literature regarding the behavior of chimpanzees and other apes in the wild to assess the potential utility of a situated-activity approach for comparative studies of primate cognition. It is argued in the Discussion that schemata constitute a common core of higher primate intelligence. Planning, emergence, and alterations of the environment to facilitate production further characterize human and chimpanzee or gorilla behaviors to varying degrees. Less apparent in the nonhuman-primate literature is evidence of governing principles, rehearsal, and skill-based reputations or identities entailing theories of mind. Nonetheless, recent observations in the wild suggest that further research is warranted to explore the rudiments of each of these components to enhance our understanding of the ecology of primate cognition and its evolutionary history.

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Pattern of maternal serum corticotropin-releasing hormone concentration during pregnancy in the common marmoset (Callithrix jacchus).

Corticotropin-releasing hormone (CRH), a potent neuropeptide, is produced by the placenta of anthropoid primates. No other mammals, including prosimian primates, are known to produce placental CRH. In humans, placental CRH appears to play an important role in the progression of pregnancy to parturition. Maternal circulating CRH begins to rise early in pregnancy and increases until parturition. Gorillas and chimpanzees share this pattern of increasing maternal CRH during pregnancy with humans. In humans, chimpanzees, and gorillas, maternal CRH and estradiol concentrations are correlated, consistent with the hypothesis that CRH is involved in the biosynthetic pathway for placental estrogen production. In contrast, in baboons, maternal circulating CRH rises precipitously early in pregnancy and then declines, though CRH is detectable until birth. This research was designed to investigate the pattern of maternal circulating CRH in the common marmoset during pregnancy. Blood samples were taken across gestation from nine subjects over 11 pregnancies, and the plasma was assayed for CRH. The pattern of maternal circulating CRH in the common marmoset was similar to that of the baboon, with a rapid rise starting at about 50 days postconception and a peak at approximately 70 days postconception. By 110 days postconception, CRH concentration had plateaued at a significantly lower value. The peak and mean values for CRH were associated with fetal number (e.g., females gestating triplets had higher values than females gestating twins). Urinary estradiol showed no association with plasma CRH concentration. Marmosets appear to differ from the great apes in this regard, and to share a pattern of maternal CRH during pregnancy with the baboon, indicating that the baboon and marmoset pattern may be ancestral. The function of the early rapid rise of CRH in baboons and marmosets, and the significance of this difference between monkeys and apes, are not known.

Animals↗

Bed-building in captive chimpanzees (Pan troglodytes): the importance of early rearing.

In the wild, great apes sleep in beds they make by successively bending branches into an interwoven platform. These beds are functionally more closely related to human beds than they are to the nests and tree-holes used by other primate species. The idea that bed-building by great apes is learned behavior that is dependent on appropriate early experiences has been proposed but never fully tested. In the present study this hypothesis was indirectly tested in 73 captive adult chimpanzees (Pan troglodytes; 27 wild-born and 46 captive-born). Bed-building and use were tested experimentally by the introduction of three sets of bedding materials. Over 200 hr of scan-sampling data were collected during 45-min observations following introduction of the bedding material. The wild-born subjects built and used beds significantly more often than the captive-born subjects. Also, wild-born subjects used more complex techniques during construction. Captive-born subjects that were mother-reared through early adolescence spent more time building and using beds than their nursery-reared counterparts. These differences remained consistent even when previous adult experience with bedding (measured as naturalistic vs. traditional housing) was accounted for. These results suggest that bed-building is a learned behavior that requires early experience and practice for acquisition.

Animals↗

Loss of N-glycolylneuraminic acid in humans: Mechanisms, consequences, and implications for hominid evolution.

The surface of all mammalian cells is covered with a dense and complex array of sugar chains, which are frequently terminated by members of a family of molecules called sialic acids. One particular sialic acid called N-glycolylneuraminic acid (Neu5Gc) is widely expressed on most mammalian tissues, but is not easily detectable on human cells. In fact, it provokes an immune response in adult humans. The human deficiency of Neu5Gc is explained by an inactivating mutation in the gene encoding CMP-N-acetylneuraminic acid hydroxylase, the rate-limiting enzyme in generating Neu5Gc in cells of other mammals. This deficiency also results in an excess of the precursor sialic acid N-acetylneuraminic acid (Neu5Ac) in humans. This mutation appears universal to modern humans, occurred sometime after our last common ancestor with the great apes, and happens to be one of the first known human-great ape genetic differences with an obvious biochemical readout. While the original selection mechanisms and major biological consequences of this human-specific mutation remain uncertain, several interesting clues are currently being pursued. First, there is evidence that the human condition can explain differences in susceptibility or resistance to certain microbial pathogens. Second, the functions of some endogenous receptors for sialic acids in the immune system may be altered by this difference. Third, despite the lack of any obvious alternate pathway for synthesis, Neu5Gc has been reported in human tumors and possibly in human fetal tissues, and traces have even been detected in normal human tissues. One possible explanation is that this represents accumulation of Neu5Gc from dietary sources of animal origin. Finally, a markedly reduced expression of hydroxylase in the brains of other mammals raises the possibility that the human-specific mutation of this enzyme could have played a role in human brain evolution.

Animals↗

Cross-site differences in foraging behavior of white-faced capuchins (Cebus capucinus).

Researchers have identified a variety of cross-site differences in the foraging behavior of free-ranging great apes, most notably among chimpanzees (Pan troglodytes) and more recently orangutans (Pongo pygmaeus), that are not due to obvious genetic or ecological differences. These differences are often referred to as "traditions." What is not known is whether this high level of interpopulation variation in behavior is limited to hominoids. In this study, we use long-term data from three Costa Rican field sites that are geographically close and similar ecologically to identify potential foraging traditions in white-faced capuchins (Cebus capucinus). Foraging traditions are predicted in Cebus because of many behavioral and morphological convergences between this genus and the great apes. The processing techniques used for the same food species were compared across sites, and all differences found were classified as present, habitual, or customary. Proximity data were also analyzed to determine if social learning mechanisms could explain variation in foraging behavior. Of the 61 foods compared, we found that 20 of them are processed differently by capuchins across sites. The differences involve pound, rub, tap, "fulcrum," "leaf-wrap," and "army ant following." For most of the differences with enough data to analyze, the average proximity score of the "matched" dyads (two individuals within a group who shared a "different" processing technique) was statistically higher than the average proximity score of the remaining "unmatched" dyads.

Animals↗

3D-kinematics of vertical climbing in hominoids.

Vertical climbing has played an important role in theories about the evolution of habitual bipedalism in early hominids and of locomotor specialization in hominoids. However, empirical data on vertical climbing in nonhuman primates are scarce, especially regarding kinematics. In this paper, the kinematics of flexed-elbow vertical climbing of four hominoid species are reported: western lowland gorillas (Gorilla gorilla gorilla), bonobos (Pan paniscus), Sumatran orangutans (Pongo pygmaeus abelii), and yellow-cheeked crested gibbons (Hylobates concolor gabriellae). The climbing sequences were recorded simultaneously by four digital video cameras in European zoos. Analyzed parameters include intersegmental flexion/extension angles of the shoulder, elbow, hip, and knee joints, horizontal abduction/adduction angles of the upper arm and thigh, and distance of the body center of gravity from the substrate. Analyses reveal that differences in climbing kinematics between individuals of different age/sex classes of one species are mostly size-related, whereas interspecific differences reflect specific locomotor adaptations. The climbing kinematics of bonobos and gorillas are more similar to each other than to those of orangutans or gibbons. The range of motion of the major limb joints in adult orangutans is larger than in African apes, although the hip is not more extended. Gibbons horizontally abduct the arm more and the thigh less during climbing than any of the great apes. These results allow us to question some qualitative descriptions of vertical climbing kinematics reported in the literature, and to discuss implications for the evolution of locomotor adaptations in hominoids.

Age Factors↗

A study of the taxonomic significance of the dental arch.

Sixteen dimensions were measured from the maxillary and mandibular dental arches of different ethnic groups of man, apes and monkeys. Multivariate analysis showed that discrimination was possible among the ethnic groups of man on the one hand and between the ape and monkey samples on the other. Nevertheless, the actual degree of discrimination between the primate samples depended upon whether the maxillary or mandibular arch dimensions were analysed. Furthermore, subsequent inclusion of the dental arch dimensions of fossil hominoid samples into the analysis confirmed the taxonomic significance of the dental arch, although its importance must await the acquisition of the more non-human primate data.

Animals↗

Relationship between tooth and long bone size.

Canonical correlations between tooth and long bone dimensions showed a greater correlation for Anglo-Saxons and apes than for Nineteenth Century Londoners, i.e., coefficients of 0.75 for gorilla, 0.72 for chimpanzee, 0.69 for orang-utans, 0.74 for Anglo-Saxons, but 0.53 for Nineteenth Century Londoners. Although based upon limited sample sizes and limited metrical profiles of teeth and long bones, the data support the thesis that modern Europeans are under reduced selection pressure to maintain tooth size compared with apes or ancient man.

Animals↗

Functional articulation of some hominoid foot bones: implications for the Olduvai (hominid 8) foot.

Previous observations on twelve fossil foot bones (Olduvai 8: Day and Napier, '64) together with multivariate morphometric studies of one of them, the talus (Day and Wood, '68) suggested human-like bipedality for this foot. Subsequent studies showed the conclusions on the talus to be wrong: The fossil talus, as defined by eight measures, does not resemble that of man but is reminiscent of those of creatures known (extant--orangutans) or believed (extinct--some fossil apes and monkeys) to be arboreal in habitus (Oxnard '72; Lisowski, et al., '74, '76). A reassessment of the entire Olduvai 8 foot is therefore necessary to answer the problem posed by a foot that has a non-human talus with, apparently, a human arched pattern of the remaining tarsal and metatarsal bones. The dry bones of a series of feet of extant hominoids have been rearticulated and are found to be close to the actual relationships presented by dissected specimens in which ligaments, articular cartilages and soft tissues are present. Similar rearticulation applied to casts of the individual Olduvai foot bones produces a structure that is not arched in the same manner as the human foot; it displays features that ally it more closely with the feet of various apes. Sections of casts of the already rearticulated Olduvai foot (from both the Wenner Gren Foundation and the Kenya National Museums) show that the human-like appearance of the original rearticulation is due to a series of incorrect osteological alignments. Although casts do not permit study of surface features, the dimensions of the casts are sufficiently accurate to permit rearticulation in this manner. It is thus clear, a) that the Olduvai foot is not adated for bipedality in the manner of man, and b) that it displays features in which it resembles the feet of arboreal creatures. Such anatomical characters as relate to bipedality in the fossil suggest usage as in an arboreal species that also walks bipedally with flattened arches (like a chimpanzee or gorilla) rather than with the high arches of man.

Adult↗

A quantitative comparison of the hominoid thalamus. IV. Posterior association nuclei-the pulvinar and lateral posterior nucleus.

Nuclear volumes, nerve cell densities, numbers of neurons, and volumes of nerve cell perikarya in the thalamic association complex, the pulvinar and lateral posterior nuclei (Pu-LP) were compared among two gibbons, one gorilla, one chimpanzee, and three humans. The human Pu has approximately twice as many neurons as do the great apes, whereas the human and gorilla LP have a similar number. The numbers of neurons in the human Pu and combined Pu-LP complex were predictable from the ape data. Nevertheless, a shift in perikaryal sizes from a unimodal to a bimodal population distinguished the human specimen. It is hypothesized that during human evolution Pu expanded in proportion to the rest of the brain, but that not all parts of Pu expanded equally.

Animals↗

The basicranium of Plio-Pleistocene hominids as an indicator of their upper respiratory systems.

Our analyses of extant primates have shown that a relationship exists between the degree of flexion of the basicranium and the location of upper respiratory structures such as the larynx and pharynx (Laitman et al., 1978). Based upon these relationships, we have previously used the basicrania of late Pleistocene hominids as a guide to the reconstruction of their upper respiratory anatomy (Laitman et al., 1979). This study continues our approach by examining the basicrania of Plio-Pleistocene hominids and reconstructing their upper respiratory systems. Nine Plio-Pleistocene hominids had basicrania complete enough to be used in this study. These included the originals of Sts 5, MLD 37/38, SK 47, SK 48, SK 83, Taung, KNM-ER 406, OH 24, and a cast of OH 5. Craniometric analysis of the basicrania of these specimens showed that they had marked similarities to those of extant pongids. These basicranial similarities between Plio-Pleistocene hominids and extant apes suggest that the upper respiratory systems of these groups were also alike in appearance. As with living nonhuman primates, the early hominids probably exhibited a larynx and pharynx positioned high in the neck. This high position would have permitted an intranarial epiglottis to be present during both normal respiration and the ingestion of a liquid bolus of food. The high position of the larynx would have also greatly restricted the supralaryngeal portion of the pharynx available to modify laryngeal sounds. It is thus possible that the Plio-Pleistocene hominids exhibited modes of breathing, swallowing and vocalizing similar to those of living apes.

Animals↗

Evolution of the lumbosacral angle.

The lumbosacral angle (LSA) was studied in 131 children ranging in age from birth to 5 years. This angle increases from an average of 20 degrees at birth to an average of 70 degrees at the age of 5 years; it remains at that level thereafter. This study demonstrates that the formation of the LSA is not related to increasing age, height, or weight. Nor do obstetrical requirements seems to play any major role in the formation of the lumbosacral angle. Rather, it appears that the development of the LSA is related to the progressive acquisition of erect posture and the ontogeny of bipedal locomotion. This angle is almost nil in the nonprimate mammals (who only infrequently stand erect). It is minimal in monkeys who occasionally assume bipedal postures and increases somewhat in living apes who engage in facultative bipedal positional behavior. In the early australopithecines, the LSA is increased over that in apes, and it reaches its maximum in Homo sapiens. Deviations from normal and healthy erect posture in Homo sapiens result in corresponding changes in the lumbosacral angle. Lumbar and sacral angles (both forming the lumbosacral angle) are almost equal in all mammalian species. Since the sacral angle of Australopithecus afarensis is approximately 15 degrees, it can be implied that its lumbosacral angle was small, thus attesting to its "imperfect" erect posture and "primitive" form of bipedal locomotion.

Adult↗

Evolutionary and morphological significance of the deflecting wrinkle in the lower molars of the Hominoidea.

The deflecting wrinkle is a well-known character state of the lower m2 and M1 of the human dentition, but there is little information regarding its presence in great apes. The deflecting wrinkle is more frequent on M2-3 in all extant pongid genera studied in this paper except Pan paniscus, in which M1 has the highest frequency (16.0%) and in which this wrinkle is absent on M3. In Gorilla, it is absent on M1, with only a low incidence on M2-3. Its greatest frequency in Pongo is always on M2 (20.2%), which is the greatest expression of the trait in the great apes. We interpret the presence of the deflecting wrinkle as an incidental effect and suggest that it represents a plesiomorphic character state in the Hominoidea.

Animals↗

Sexual dimorphism in Laccopithecus robustus, a late miocene hominoid from China.

Laccopithecus robustus is a siamang-sized fossil ape from the Miocene site of Lufeng, China. The species is known from a partial cranium, numerous mandibles, and scores of isolated teeth. This species shows striking dental similarities to Pliopithecus from the Miocene of Europe and a number of cranial similarities to extant gibbons. Laccopithecus differs from extant gibbons and resembles other fossil and extant apes in showing marked sexual dimorphism in the size and shape of the canines and anterior lower premolars. Evidence for sexual differences in either the size or shape of other teeth is less clear. There is some evidence for a sexual size dimorphism based on the variability of molar teeth.

Animals↗

EMG of serratus anterior and trapezius in the chimpanzee: scapular rotators revisited.

The importance of arm-raising has been a major consideration in the functional interpretation of differences in shoulder morphology among species of nonhuman primates. Among the characters that have been associated with enhancement of the arm-raising mechanism in hominoid primates are the relative enlargement of cranial trapezius and caudal serratus anterior, as the main scapular rotators, as well as changes in scapular morphology associated with their improved leverage for scapular rotation. Yet in an EMG study of cranial trapezius and caudal serratus anterior function in the great apes, Tuttle and Basmajian (Yrbk. Phys. Anthropol. 20:491-497, 1977) found these muscles to be essentially inactive during arm-raising. Although Tuttle and Basmajian suggest that the cranial orientation of the glenoid fossa in apes has reduced the demand for scapular rotation during arm-raising, subsequent EMG studies on other primate species suggest that these muscles do play a significant role in arm motion during active locomotion. This paper presents a reexamination of muscle recruitment patterns for trapezius and caudal serratus anterior in the chimpanzee. All but the lowest parts of caudal serratus anterior were found to be highly active during arm-raising motions, justifying earlier morphological interpretations of differences in caudal serratus anterior development. The lowest digitations of this muscle, while inactive during arm-raising, displayed significant activity during suspensory postures and locomotion, presumably to control the tendency of the scapula to shift cranially relative to the rib cage. Cranial trapezius did not appear to be involved in arm-raising; instead, its recruitment was closely tied to head position.

Animals↗