Human Adaptation, Somatotyping and Demography: Bergmann's and Allen's Rules, the Human Adaptability Programme, Adaptation to Heat, Cold and High Altitude, Somatotyping from Kretschmer and Sheldon to Parnell and Heath–Carter, and Demography — Its Multidisciplinary Nature, Anthropological Demography, Fertility, Morbidity, Mortality and Migration, and Crow's Index of Selection Intensity Worked

The second half of Unit IV takes three topics that share a concern with the whole body in its environment. Human adaptation asks how populations cope with heat, cold and thin air, through the ecogeographic rules of Bergmann and Allen, the international Human Adaptability Programme and the physiology of acclimatisation and genetic adaptation. Somatotyping asks how body build varies and how it is scored, from Kretschmer's psychiatric types and Sheldon's three components to Parnell's and Heath and Carter's anthropometric methods — the syllabus prints "Parnoll" and "Health-Carter", which are Parnell and Heath–Carter. Demography asks how populations grow and change, through fertility, morbidity, mortality and migration, and how anthropological demography differs from the demographer's own; Crow's index of the opportunity for selection connects it back to evolution. NET Physical Education teaches somatotyping for the coach and NET Economics demography for the planner; here both are the biological anthropologist's tools, and the rates and the index are worked with checked numbers.

1. Bergmann's and Allen's rules and the Human Adaptability Programme

Adaptation is any feature — genetic, developmental, physiological or cultural — that improves an organism's fit to its environment. Anthropologists distinguish genetic adaptation (selected over generations: sickle cell against malaria, the Tibetan EPAS1 allele), developmental adaptation or plasticity (acquired irreversibly during growth: the large chests of Andean children), acclimatisation (reversible physiological adjustment within a lifetime: sweating more after weeks in the heat) and cultural adaptation (clothing, housing, fire, diet), which in humans buffers all the rest. Two ecogeographic rules, framed for warm-blooded animals, apply to human body form. Bergmann's rule (Carl Bergmann, 1847): within a species, populations in colder climates have larger body mass, because volume (heat production) rises faster than surface area (heat loss) as size increases — so a large body conserves heat and a small one sheds it; Roberts (1953) showed mean body weight in humans falls with mean annual temperature. Allen's rule (Joel Allen, 1877): populations in colder climates have shorter extremities — limbs, ears, noses — relative to the trunk, again reducing surface area per unit mass; the stocky, short-limbed Inuit and the linear, long-limbed Nilotes are the textbook pair, and the Neanderthal body is the fossil case. Both rules are read through the surface-to-volume ratio and the relative sitting height or the crural index, and both are statistical tendencies with exceptions produced by diet, activity and gene flow. The Human Adaptability Programme (HAP) was one of the seven sections of the International Biological Programme (IBP, 1964–74), directed on the human side by Joseph Weiner: a coordinated, worldwide study of human populations in extreme environments — high altitude (Andes, Himalaya, Ethiopia), the Arctic (Inuit, Sami), the tropics and deserts (Kalahari San, Australian Aborigines), and isolated islands — using standardised methods of anthropometry, physiology, nutrition, genetics and demography (Weiner and Lourie's IBP Handbook No. 9, 1969). It established the comparative baseline for everything in this chapter, and Indian participation (the Anthropological Survey, the universities) produced data on tribal and high-altitude groups.

⚠️ Bergmann is mass, Allen is limbs
Both rules say cold → conserve heat by lowering surface area per unit volume. Bergmann does it with a larger body; Allen with shorter appendages. Gloger's rule (darker pigmentation in humid warm climates) is the third ecogeographic rule sometimes added. The HAP is part of the IBP (1964–74), not of the Human Genome Project or WHO.

2. Adaptation to heat, cold and high altitude

StressShort-term responsesAcclimatisation and developmental adaptationGenetic adaptation and population examples
Heat (dry desert; humid tropics)Vasodilation of the skin; sweating (humans have 2–4 million eccrine glands, the most of any mammal); reduced activity; heat exhaustion and heat stroke if it failsAfter 1–2 weeks: earlier and heavier sweating with less salt, expanded plasma volume, lower heart rate; humid heat is harder because sweat cannot evaporateLinear, long-limbed build (Allen) and low mass (Bergmann) in the Nilotes and other tropical groups; dark skin protecting folate and against UV damage; cultural: loose clothing, siesta, evaporative cooling
ColdVasoconstriction; shivering; piloerection (useless in humans); cold-induced vasodilation in the hands ("hunting response", Lewis, 1930)Habituation: less shivering and less discomfort; raised basal metabolic rate (Inuit ~15–30% above standard, partly diet); non-shivering thermogenesis via brown fat in infantsStocky, short-limbed build (Bergmann, Allen) in Arctic peoples; Inuit hands with high blood flow; Australian Aborigines sleep through cold nights by letting skin temperature fall (insulative response); Alacaluf of Tierra del Fuego with a raised metabolic response; cultural: fur clothing, the igloo, fire
High altitude (hypoxia above ~2,500 m; also cold, UV, low humidity)Faster, deeper breathing (hyperventilation); raised heart rate; acute mountain sickness; pulmonary and cerebral oedema in the worst casesOver days to weeks: more red cells and haemoglobin (via erythropoietin), a right-shifted oxygen dissociation curve (2,3-DPG), more capillaries; children growing at altitude develop large chests and lung volumes (developmental adaptation)Andean highlanders (Quechua, Aymara): high haemoglobin, barrel chest; Tibetans: normal haemoglobin but higher breathing rate and blood flow, the EPAS1 and EGLN1 alleles (EPAS1 of Denisovan origin), lower birth-weight loss; Ethiopian Amhara: a third pattern; chronic mountain sickness (Monge's disease) where adaptation fails
ℹ️ Two ways up the mountain
The Andean solution is more haemoglobin per litre of blood; the Tibetan solution, older by some millennia, is more air and more blood flow with ordinary haemoglobin, driven by EPAS1 (inherited from Denisovans, 2014) and EGLN1. Beall's comparison of the two is the standard citation, and the question "which population does NOT show elevated haemoglobin at altitude" wants Tibetans.

3. Somatotyping: Kretschmer, Sheldon, Parnell and Heath–Carter, and its applications

Somatotyping is the classification and quantification of body build (physique). Ernst Kretschmer, a German psychiatrist, in Körperbau und Charakter (Physique and Character, 1921), sorted patients into the pyknic (round, stocky, short-necked — associated in his sample with manic-depressive illness), the athletic (muscular, broad-shouldered), the leptosomic or asthenic (thin, narrow, long-limbed — associated with schizophrenia) and the dysplastic (mixed or abnormal); his typology and his psychiatric correlations are historical. William Sheldon, in The Varieties of Human Physique (1940, with Stevens and Tucker), replaced types with three continuous components named for the embryonic germ layers and rated each on a seven-point scale from standardised photographs (photoscopic method) with height/∛weight as the guide: endomorphy (roundness, softness, predominance of the digestive viscera), mesomorphy (muscle and bone, the athletic build) and ectomorphy (linearity, fragility, a large surface relative to mass). A somatotype is written as three digits in that order — 7-1-1 the extreme endomorph, 1-7-1 the extreme mesomorph, 1-1-7 the extreme ectomorph, 4-4-4 balanced — and Sheldon's claim of correlated temperaments (viscerotonia, somatotonia, cerebrotonia) in The Varieties of Temperament (1942) is discarded. R.W. Parnell (Behaviour and Physique, 1958) made the rating anthropometric rather than photographic, with the M.4 deviation chart: fat (three skinfolds), muscularity (bone breadths and limb girths) and linearity (height/∛weight) scored against age-specific tables, keeping Sheldon's 1–7 scale. Barbara Heath and Lindsay Carter (1967; Carter and Heath, Somatotyping: Development and Applications, 1990) produced the method now standard: an anthropometric somatotype from ten measurements — height, weight, four skinfolds (triceps, subscapular, supraspinale, medial calf), two bone breadths (biepicondylar humerus and femur) and two girths (flexed upper arm and calf) — with an open-ended rating scale (not capped at 7), age- and sex-independent, and a somatochart on which the three components are plotted as a single point by the coordinates x = ectomorphy − endomorphy and y = 2 × mesomorphy − (endomorphy + ectomorphy). Applications: kinanthropometry and sports science (the mesomorphic profiles of wrestlers and sprinters, the ecto-mesomorphy of high-jumpers, the endo-mesomorphy of shot-putters), growth and nutrition studies (somatotype changes with age and undernutrition), ergonomics, medicine (the association of endomorphy with cardiovascular risk), and the anthropological comparison of populations — Indian tribal and caste groups have been somatotyped in a large literature from the 1970s.

SystemYearBasisCategories or scaleNote
Kretschmer1921Clinical observation of psychiatric patientsPyknic, athletic, leptosomic (asthenic), dysplasticTypology; the psychiatric correlations are not upheld
Sheldon1940Photoscopic rating of standardised photographs; height/∛weightEndomorphy, mesomorphy, ectomorphy, each 1–7; three-digit somatotypeFirst continuous system; temperament claims discarded
Parnell ("Parnoll" in the syllabus)1958Anthropometric: skinfolds, bone breadths, girths, height/∛weight; M.4 deviation chartFat, muscularity, linearity on Sheldon's 1–7 scaleMade the rating objective and repeatable
Heath–Carter ("Health-Carter" in the syllabus)1967Ten anthropometric measurements; open-ended scale; somatochartEndomorphy, mesomorphy, ectomorphy without an upper limit; any age and sexThe current international standard (ISAK)
🧠 Endo, meso, ecto — in that order, always
A somatotype is written endomorphy-mesomorphy-ectomorphy, so 2-6-2 is a strong mesomorph and 5-3-1 an endomorph. Sheldon's names come from the germ layers (endoderm → gut, mesoderm → muscle and bone, ectoderm → skin and nervous system). Kretschmer's pyknic ≈ endomorph, athletic ≈ mesomorph, leptosomic ≈ ectomorph, but Sheldon's are continuous components, not types.

4. Demography: its multidisciplinary nature, anthropological demography, and the measures of fertility, morbidity, mortality and migration, worked

Demography (Achille Guillard coined the word in 1855) is the statistical study of human populations — their size, composition and distribution and the three processes that change them: fertility, mortality and migration. It is multidisciplinary by nature: its data come from censuses, vital registration and surveys (statistics); its explanations from economics (Malthus's Essay of 1798; the demographic transition of Thompson 1929 and Notestein 1945, from high to low birth and death rates), sociology (family, marriage, education), geography (distribution, urbanisation), public health and medicine (morbidity, mortality), biology and genetics (fecundity, selection) and history (reconstitution from parish registers). Anthropological demography is the anthropologist's contribution and complement: it studies small populations — a tribe, a caste, a village, a band — often without registration data, by genealogies, censuses of its own and long residence; it treats fertility, marriage and death as embedded in kinship, ritual and economy rather than as individual choices; it reads the demography of hunter-gatherers (Howell's Dobe !Kung, 1979) and of pastoralists and peasants as adaptations; and it feeds population genetics with the mating structure and reproductive variance that selection works on. Its measures are the demographer's. Fertility (actual reproduction, as against fecundity, the biological capacity) is measured by the crude birth rate (live births per 1,000 mid-year population), the general fertility rate (births per 1,000 women aged 15–49), age-specific fertility rates and the total fertility rate (the average number of children a woman would bear over her life at current age-specific rates — replacement level about 2.1), with the gross and net reproduction rates counting daughters; its determinants are the proximate ones of Davis and Blake (1956) and Bongaarts (1978) — marriage, contraception, abortion, breastfeeding and postpartum abstinence, sterility — behind which stand education, income, religion, infant mortality and the value of children. Morbidity, the burden of illness, is measured by incidence (new cases per 1,000 per year) and prevalence (existing cases at a point of time), and its determinants are nutrition, sanitation, crowding, occupation and health services. Mortality is measured by the crude death rate (deaths per 1,000 mid-year population), age- and cause-specific rates, the infant mortality rate (deaths under one year per 1,000 live births, the most sensitive index of living conditions), the under-five and maternal mortality rates, and life expectancy at birth from the life table; its determinants are nutrition, infection, sanitation, medical care, income, sex (women outlive men) and age. Migration is a change of usual residence across a boundary, internal or international, in-migration and out-migration giving net migration; Ravenstein's laws (1885), Lee's push and pull factors (1966), Zelinsky's mobility transition, and the anthropological types — seasonal, circular, chain and forced migration, rural–urban and tribal displacement — are its frame, and its determinants are employment, land, marriage, education, conflict and disaster. Worked: a district has a mid-year population of 50,000; in the year there were 1,500 live births, 400 deaths and 60 infant deaths. CBR = 1,500 ÷ 50,000 × 1,000 = 30 per 1,000; CDR = 400 ÷ 50,000 × 1,000 = 8 per 1,000; the rate of natural increase = 30 − 8 = 22 per 1,000 = 2.2% a year; IMR = 60 ÷ 1,500 × 1,000 = 40 per 1,000 live births. If the seven five-year age-specific fertility rates (15–19 to 45–49) sum to 700 per 1,000 women, TFR = 5 × 700 ÷ 1,000 = 3.5 children per woman.

⚠️ Denominators
CBR and CDR are per 1,000 of the whole mid-year population; IMR is per 1,000 live births, not per 1,000 population; GFR is per 1,000 women aged 15–49; TFR is a sum of age-specific rates (× 5 for five-year groups), not a rate per 1,000. Fertility is performance, fecundity is capacity. The distractor puts infant deaths over total population.

5. Selection intensity: Crow's index, worked

Natural selection can act only where individuals differ in survival to reproductive age and in the number of offspring they leave, and James Crow (1958) proposed an index of the total opportunity for selection that a population's demography allows, computed from those two components. The mortality component Im = Pd ÷ Ps, where Pd is the proportion of live-born who die before reproductive age and Ps = 1 − Pd the proportion who survive. The fertility component If = Vf ÷ x̄², where x̄ is the mean number of live births per surviving woman (or per person) who reaches reproductive age and Vf the variance of that number. The total index I = Im + If ÷ Ps — the fertility component is divided by Ps because only survivors reproduce. Worked: in a tribal population 20% of children die before maturity, so Pd = 0.2, Ps = 0.8 and Im = 0.2 ÷ 0.8 = 0.25. Among surviving women the mean number of live births is 4 with a variance of 4, so If = 4 ÷ 16 = 0.25. I = 0.25 + 0.25 ÷ 0.8 = 0.25 + 0.3125 = 0.5625. If mortality is later halved (Pd = 0.1: Im = 0.111) and the variance of fertility stays, I = 0.111 + 0.25 ÷ 0.9 = 0.111 + 0.278 = 0.389: the index falls, and the fertility component now supplies most of the opportunity for selection — the pattern Crow and Spuhler found as populations pass through the demographic transition. The index measures opportunity, not the direction of selection or its genetic effect; it has been computed for many Indian tribal and caste populations (Basu, Reddy, Bhasin and others), with typical values between 0.3 and 1.5, higher where child mortality and variance in family size are high. Selection intensity so measured is the link between the demographer's rates and the evolutionist's forces, which is why the syllabus places it here.

🧠 Crow in three lines
Im = Pd/Ps (deaths before reproduction, over survivors). If = Vf/x̄² (variance of family size over the square of its mean). I = Im + If/Ps. With Pd = 0.2, x̄ = 4, Vf = 4: 0.25 + 0.25/0.8 = 0.5625. Forgetting to divide If by Ps (giving 0.50) is the planted error.

Key takeaways

  • Adaptation is genetic, developmental, physiological (acclimatisation) or cultural; Bergmann's rule (1847) says colder climates favour larger body mass, Allen's rule (1877) shorter extremities — both lower surface area per unit volume; the Human Adaptability Programme was the human section of the International Biological Programme (1964–74), led by Weiner.
  • Heat: vasodilation, sweating (2–4 million eccrine glands), then heavier dilute sweat and plasma expansion; cold: vasoconstriction, shivering, the hunting response, raised metabolism (Inuit), insulative sleep (Aborigines); altitude: hyperventilation, then more haemoglobin (Andeans) or more ventilation and blood flow with the EPAS1 allele (Tibetans), and large chests from developmental adaptation.
  • Somatotyping: Kretschmer (1921: pyknic, athletic, leptosomic, dysplastic), Sheldon (1940: endomorphy, mesomorphy, ectomorphy rated 1–7 from photographs; 7-1-1, 1-7-1, 1-1-7), Parnell (1958: anthropometric M.4 chart) and Heath–Carter (1967: ten measurements, open scale, somatochart — the standard); the syllabus's "Parnoll" and "Health-Carter" are Parnell and Heath–Carter.
  • Demography (Guillard 1855) is multidisciplinary; anthropological demography studies small populations through genealogies and treats vital events as embedded in kinship and economy; CBR and CDR per 1,000 population, IMR per 1,000 live births, TFR = 5 × Σ ASFR ÷ 1,000 (replacement ~2.1) — a district with 50,000 people, 1,500 births, 400 deaths and 60 infant deaths has CBR 30, CDR 8, natural increase 2.2% and IMR 40.
  • Crow's index of the opportunity for selection (1958): Im = Pd/Ps, If = Vf/x̄², I = Im + If/Ps — with 20% pre-reproductive mortality and a mean of 4 births with variance 4, I = 0.25 + 0.3125 = 0.5625; the index falls and shifts toward the fertility component as mortality declines.

Practice questions (10)

Attempt each one before opening the answer. Every explanation names the tempting wrong option as well as the right one, because that is where marks are lost.

  1. The rule that populations of a warm-blooded species living in colder climates tend to have larger body mass is

    1. Gloger's rule
    2. Bergmann's rule
    3. Cope's rule
    4. Allen's rule
    Show answer

    Answer: B — Bergmann's rule

    Bergmann (1847) related body mass to climate through the surface-to-volume ratio; Allen (1877) related the length of the extremities; Gloger concerns pigmentation and Cope the tendency of lineages to grow larger over evolutionary time.
  2. In Sheldon's system a somatotype written 1-7-1 describes

    1. an extreme endomorph — round and soft
    2. an extreme mesomorph — muscular and heavy-boned
    3. a balanced physique
    4. an extreme ectomorph — linear and fragile
    Show answer

    Answer: B — an extreme mesomorph — muscular and heavy-boned

    The three digits are always endomorphy, mesomorphy and ectomorphy in that order, each rated 1–7; 7-1-1 is the extreme endomorph, 1-1-7 the extreme ectomorph and 4-4-4 balanced.
  3. A district with a mid-year population of 50,000 recorded 1,500 live births in a year, of which 60 children died before their first birthday. What is the infant mortality rate per 1,000 live births?

    Numerical answer — type the value.

    Show answer

    Answer: 40

    IMR = infant deaths ÷ live births × 1,000 = 60 ÷ 1,500 × 1,000 = 40. The denominator is live births, not population; dividing by 50,000 would give the meaningless 1.2.
  4. Match the somatotyping system with its distinctive feature. (a) Kretschmer (b) Sheldon (c) Parnell (d) Heath–Carter. Features: (1) Ten anthropometric measurements and an open-ended scale (2) Pyknic, athletic and leptosomic types from psychiatric practice (3) Photoscopic rating of three components on a 1–7 scale (4) The M.4 deviation chart of fat, muscularity and linearity

    1. a-2, b-3, c-1, d-4
    2. a-2, b-4, c-3, d-1
    3. a-2, b-3, c-4, d-1
    4. a-3, b-2, c-4, d-1
    Show answer

    Answer: C — a-2, b-3, c-4, d-1

    Kretschmer (1921) gave the psychiatric typology, Sheldon (1940) the photoscopic three-component rating, Parnell (1958) the anthropometric M.4 chart and Heath–Carter (1967) the ten-measurement open-scale method that is now standard.
  5. Tibetan highlanders differ from Andean highlanders in their adaptation to hypoxia chiefly in that Tibetans

    1. show no physiological adaptation at all
    2. rely entirely on cultural adaptation such as clothing
    3. maintain near-normal haemoglobin but breathe faster and have higher blood flow, associated with the EPAS1 and EGLN1 alleles
    4. have markedly raised haemoglobin concentrations and larger chests
    Show answer

    Answer: C — maintain near-normal haemoglobin but breathe faster and have higher blood flow, associated with the EPAS1 and EGLN1 alleles

    Beall's comparison showed the Andean pattern of high haemoglobin and the Tibetan pattern of high ventilation and blood flow with ordinary haemoglobin; the Tibetan EPAS1 variant, inherited from Denisovans, is the best-known genetic adaptation in humans.
  6. Which of the following are proximate determinants of fertility in the Davis–Blake and Bongaarts frameworks? Select all that apply.

    1. Contraceptive use
    2. Proportion of women married
    3. Duration of breastfeeding and postpartum infecundability
    4. Female literacy
    Show answer

    Answer: A — Contraceptive use; B — Proportion of women married; C — Duration of breastfeeding and postpartum infecundability

    Marriage, contraception, abortion and lactational infecundability are the proximate (biological and behavioural) determinants through which everything else acts; literacy, income and religion are background or distal determinants.
  7. In a population 20% of live-born children die before reproductive age; among surviving women the mean number of live births is 4 with a variance of 4. What is Crow's index of total selection intensity? Give the answer to four decimal places.

    Numerical answer — type the value.

    Show answer

    Answer: 0.5625

    Im = Pd/Ps = 0.2/0.8 = 0.25; If = Vf/x̄² = 4/16 = 0.25; I = Im + If/Ps = 0.25 + 0.25/0.8 = 0.25 + 0.3125 = 0.5625. Omitting the division of If by Ps gives the wrong 0.50.
  8. The seven five-year age-specific fertility rates of a population, for women aged 15–19 to 45–49, sum to 700 births per 1,000 women. The total fertility rate is

    1. 700 per 1,000 women
    2. 7.0 children per woman
    3. 0.7 children per woman
    4. 3.5 children per woman
    Show answer

    Answer: D — 3.5 children per woman

    Each five-year rate applies for five years of a woman's life, so TFR = 5 × 700 ÷ 1,000 = 3.5, well above the replacement level of about 2.1.
  9. Assertion (A): As a population passes through the demographic transition, Crow's index of selection intensity falls and its fertility component comes to dominate. Reason (R): Declining pre-reproductive mortality leaves the mortality component unchanged, because that component is measured by the variance in family size.

    1. Both A and R are true, and R is the correct explanation of A
    2. Both A and R are true, but R is not the correct explanation of A
    3. A is true, but R is false
    4. A is false, but R is true
    Show answer

    Answer: C — A is true, but R is false

    A is true: in the worked example halving mortality lowered I from 0.5625 to 0.389 and raised the fertility component's share from 56% to 71%, the pattern Crow and Spuhler described. R is false: the mortality component is Im = Pd/Ps, which falls as the proportion dying before reproductive age falls; the variance in family size belongs to the fertility component, If = Vf/x̄².
  10. Which statements about human adaptation to cold are correct? Select all that apply.

    1. A stocky build with short limbs, as among the Inuit, follows Bergmann's and Allen's rules
    2. Australian Aborigines sleeping in the cold show an insulative response, letting skin temperature fall without raising metabolism
    3. The "hunting response" is a periodic cold-induced vasodilation of the hands that protects them from frostbite
    4. Piloerection is an effective source of insulation in humans
    Show answer

    Answer: A — A stocky build with short limbs, as among the Inuit, follows Bergmann's and Allen's rules; B — Australian Aborigines sleeping in the cold show an insulative response, letting skin temperature fall without raising metabolism; C — The "hunting response" is a periodic cold-induced vasodilation of the hands that protects them from frostbite

    Piloerection (goose-flesh) is a vestigial reflex that traps no useful air in the sparse human coat; the hunting response, the Aboriginal insulative pattern and the Arctic body build are correctly stated.