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Korean Journal of Psychology: General

Psychological Significance of Finger Length Ratio: 2D:4D as a Marker of Prenatal Testosterone

Korean Journal of Psychology: General / Korean Journal of Psychology: General, (P)1229-067X; (E)2734-1127
2014, v.33 no.4, pp.787-814


Abstract

It has been known that testosterone in utero is involved in structural and functional organization of the brain and has permanent effects on later behavior. The same hormone that affects early brain development also contributes to the establishment of finger length patterns. 2D:4D is a proxy biomarker of prenatal testosterone level, as low 2D:4D reflects high testosterone relative to estrogen, and a sexually dimorphic trait, as males tend to have lower 2D:4D than females. The aim of this review is to provide the physiological evidence that 2D:4D ratio can be used as a putative biomarker of prenatal testosterone levels and to give supporting evidence. In addition, this study examined the relationships between 2D:4D and sex-hormone-mediated traits (psychological, physical, and behavioral traits). Atypical levels of prenatal testosterone may be the cause of atypical behaviors and can be a possible indicator of a breakdown of developmental homeostatis, which may predispose individuals to certain diseases. 2D:4D was also associated with body shape and size, handgrip strength, and physical prowess. Therefore, 2D:4D is likely to be indirect biological indicator of prenatal testosterone levels that can be used to examine the effects of prenatal testosterone on various behaviors.

keywords
2D:4D, 태내 테스토스테론, 대뇌 편측성, 신체적 특징, 사회적 행동, 2D:4D, prenatal testosterone, cerebral laterality, physical traits, social behavior

Reference

1.

Archer, J. (2004). Sex differences in aggression in real-world settings: A meta-analytic review. Review of General Psychology, 8, 291-322.

2.

Arnold, A. P., & Breedlove, S. M. (1985). Organizational and activational effects of sex steroids on brain and behavior: a reanalysis. Hormones and Behavior, 19, 469-498.

3.

Austin, E. J., Manning, J. T., McInroy, K., & Mathews, E. (2002). A preliminary investigation of the associations between personality, cognitive ability and digit ratio. Personality and Individual Differences, 33, 1115-1124.

4.

Auyeung, B., Taylor, K., Hackett, G., & Baron-Cohen, S. (2010). Research Foetal testosterone and autistic traits in 18 to 24-month-old children. Molecular Autism, 1, 1-8.

5.

Bailey, A. A., & Hurd, P. L. (2005a). Finger length ratio (2D:4D) correlates with physical aggression in men but not in women. Biological psychology, 68, 215-222.

6.

Bailey, A. A., & Hurd, P. L. (2005b). Depression in men is associated with more feminine finger length ratios. Personality and Individual Differences, 39, 829-836.

7.

Barker, D. J. P. & Sultan, H. Y. (1995). Fetal programming of human disease. In M. A. Hanson, J. A. D. Spencer, & C. H. Rodeck (Eds.), Growth (pp.255-276). Cambridge, MA:Cambridge University Press.

8.

Baron-Cohen, S. (2002). The extreme male brain theory of autism. Trends in Cognitive Sciences, 6, 248-254.

9.

Benderlioglu, Z., & Nelson, R. J. (2004). Digit length ratios predict reactive aggression in women, but not in men. Hormones and Behavior, 46, 558-564.

10.

Ben-Hur, H., Thole, H. H., Mashiah, A., Insler, V., Berman, V., Shezen, E., ... & Ornoy, A. (1997). Estrogen, progesterone and testosterone receptors in human fetal cartilaginous tissue:immunohistochemical studies. Calcified Tissue International, 60, 520-526.

11.

Benderlioglu, Z., & Nelson, R. J. (2004). Digit length ratios predict reactive aggression in women, but not in men. Hormones and Behavior, 46, 558-564.

12.

Bennett, M., Manning, J. T., Cook, C. J., & Kilduff, L. P. (2010). Digit ratio (2D:4D) and performance in elite rugby players. Journal of Sports Sciences, 28, 1415-1421.

13.

Berenbaum, S. A., Bryk, K. K., Nowak, N., Quigley, C. A., & Moffat, S. (2009). Fingers as a marker of prenatal androgen exposure. Endocrinology, 150, 5119-5124.

14.

Berenbaum, S. A., & Resnick, S. M. (1997). Early androgen effects on aggression in children and adults with congenital adrenal hyperplasia. Psychoneuroendocrinology, 22, 505-515.

15.

Bescos, R., Esteve, M., Porta, J., Mateu, M., Irurtia, A., & Voracek, M. (2009). Prenatal programming of sporting success: Associations of digit ratio (2D:4D), a putative marker for prenatal androgen action, with world rankings in female fencers. Journal of Sports Sciences, 27, 625-632.

16.

Bjorntorp, P. (1987). Fat cell distribution and metabolism. In R. J. Wurtman & J. J. Wurtman (Eds.), Human Obesity (pp.66-72). NewYork: New York Academy of Sciences.

17.

Bhasin, S., Woodhouse, L., & Storer, T. W. (2001). Hormones and sport. Proof of the effect of testosterone on skeletal muscle. Journal of Endocrinology, 170, 27-38.

18.

Breedlove, S. M. (1994). Sexual differentiation of the human nervous system. Annual Review of Psychology, 45, 389-418.

19.

Breedlove, S. M. (2010). Minireview: Organizational hypothesis: instances of the fingerpost. Endocrinology, 151, 4116-4122.

20.

Brown, W. M., Hines, M., Fane, B. A., & Breedlove, S. M. (2002). Masculinized finger length patterns in human males and females with congenital adrenal hyperplasia. Hormones and Behavior, 42, 380-386.

21.

Buck, J. J., Williams, R. M., Hughes, I. A., & Acerini, C. L. (2003). In‐utero androgen exposure and 2nd to 4th digit length ratio—comparisons between healthy controls and females with classical congenital adrenal hyperplasia. Human Reproduction, 18, 976-979.

22.

Butovskaya, M. L., Vasilyev, V. A., Lazebny, O. E., Burkova, V. N., Kulikov, A. M., Mabulla, A., ... & Ryskov, A. P. (2012). Aggression, digit ratio, and variation in the androgen receptor, serotonin transporter, and dopamine D4 receptor genes in African Foragers: The Hadza. Behavior Genetics, 42, 647-662.

23.

Chamberlain, N. L., Driver, E. D., & Miesfeld, R. L. (1994). The length and location of CAG trinucleotide repeats in the androgen receptor N-terminal domain affect transactivation function. Nucleic Acids Research, 22, 3181-3186.

24.

Chapman, E., Baron-Cohen, S., Auyeung, B., Knickmeyer, R., Taylor, K., & Hackett, G. (2006). Fetal testosterone and empathy:Evidence from the empathy quotient (EQ) and the “reading the mind in the eyes” test. Social Neuroscience, 1, 135-148.

25.

Chatterjee, S., & Chowdhuri, B. J. (1991). Comparison of grip strength and isometric endurance between the right and left hands of men and their relationship with age and other physical parameters. Journal of Human Ergology, 20, 41-50.

26.

Coates, J. M., Gurnell, M., & Rustichini, A. (2009). Second-to-fourth digit ratio predicts success among high-frequency financial traders. Proceedings of the National Academy of Sciences, 106, 623-628.

27.

Cohen-Bendahan, C. C., van de Beek, C., & Berenbaum, S. A. (2005). Prenatal sex hormone effects on child and adult sex-typed behavior: methods and findings. Neuroscience &Biobehavioral Reviews, 29, 353-384.

28.

Coyne, S. M., Manning, J. T., Ringer, L., & Bailey, L. (2007). Directional asymmetry (right-left differences) in digit ratio (2D:4D)predict indirect aggression in women. Personality and Individual Differences, 43, 865-872.

29.

Crewther, B. T., Cook, C., Cardinale, M., Weatherby, R. P., & Lowe, T. (2011). Two emerging concepts for elite athletes. Sports Medicine, 41, 103-123.

30.

Csatho, A., Osvath, A., Karadi, K., Bicsak, E., Manning, J., & Kallai, J. (2003). Spatial navigation related to the second to fourth digit ratio in women. Learning and Individual Difference, 13, 239-249.

31.

Dabbs, J. M., & Dabbs, M. G. (2000). Heroes, rogues, and lovers: Testosterone and behavior. McGraw-Hill.

32.

Dadds, M. R., Allen, J. L., Oliver, B. R., Faulkner, N., Legge, K., Moul, C., ... & Scott, S. (2012). Love, eye contact and the developmental origins of empathy v. psychopathy. The British Journal of Psychiatry, 200, 191-196.

33.

De Bruin, E. I., De Nijs, P. F., Verheij, F., Verhagen, D. H., & Ferdinand, R. F. (2009). Autistic features in girls from a psychiatric sample are strongly associated with a low 2D:4D ratio. Autism, 13, 511-521.

34.

Ecker, A. (1878). Einige Bemerkungen über einen schwankenden Charakter in der Hand des Menschen [Some remarks on a varying property of the human hand]. Archiv für Anthropologie (Braunschweig), 8, 67-74.

35.

Fairburn, C. G., & Harrison, P. J. (2003). Eating disorders. The Lancet, 361, 407-416.

36.

Feliciano, P., Niemitz, E., & Vogan, K. (2011). Sexually dimorphic digits. Nature Genetics, 43, 928.

37.

Fink, B., Manning, J. T., & Neave, N. (2005). The 2nd-4th digit ratio (2D:4D) and neck circumference: implications for risk factors in coronary heart disease. International Journal of Obesity, 30, 711-714.

38.

Fink, B., Manning, J. T., Neave, N., & Tan, U. (2004). Second to fourth digit ratio and hand skill in Austrian children. Biological Psychology, 67, 375-384.

39.

Fink, B., Neave, N., & Manning, J. T. (2003). Second to fourth digit ratio, body mass index, waist-to-hip ratio, and waist-to-chest ratio:their relationships in heterosexual men and women. Annals of Human Biology, 30, 728-738.

40.

Fink, B., Thanzami, V., Seydel, H., & Manning, J. T. (2006). Digit ratio and hand-grip strength in German and Mizos men: cross-cultural evidence for an organizing effect of prenatal testosterone on strength. American Journal of Human Biology, 18, 776-782.

41.

Foundas, A. L., Leonard, C. M., & Heilman, K. M. (1995). Morphologic cerebral asymmetries and handedness. The pars triangularis and planum temporale. Archives of Neurology, 52, 501-508.

42.

Frederick, M. J. (2010). Effects of early developmental stress on adult physiology and behavior. (Doctoral dissertation, the University at Albany, State University of New York).

43.

Gallup, A. C., White, D. D., & Gallup, G. G. (2007). Handgrip strength predicts sexual behavior, body morphology, and aggression in male college students. Evolution and Human Behavior, 28, 423-429.

44.

Gann, P. H., Hennekens, C. H., Ma, J., Longcope, C., & Stampfer, M. J. (1996). Prospective study of sex hormone levels and risk of prostate cancer. Journal of the National Cancer Institute, 88, 1118-1126.

45.

Garbarino, E., Slonim, R., & Sydnor, J. (2011). Digit ratios (2D:4D) as predictors of risky decision making for both sexes. Journal of Risk and Uncertainty, 42, 1-26.

46.

Gaub, M., & Carlson, C. L. (1997). Gender differences in ADHD: a meta-analysis and critical review. Journal of the American Academy of Child & Adolescent Psychiatry, 36, 1036-1045.

47.

Geschwind, N., & Galaburda, A. M. (1985). Cerebral lateralization: Biological mechanisms, associations, and pathology: I. A hypothesis and a program for research. Archives of Neurology, 42, 428-459.

48.

Grimshaw, G. M., Bryden, M. P., & Finegan, J. K. (1995). Relations between prenatal testosterone and cerebral lateralization in children. Neuropsychology, 9, 68-79.

49.

Gur, R. C., Richard, J., Calkins, M. E., Chiavacci, R., Hansen, J. A., Bilker, W. B., ... & Gur, R. E. (2012). Age group and sex differences in performance on a computerized neurocognitive battery in children age 8-21. Neuropsychology, 26, 251.

50.

Hampson, E., Ellis, C. L., & Tenk, C. M. (2008). On the relation between 2D:4D and sex dimorphic personality traits. Archives of Sexual Behavior, 37, 133-144.

51.

Hampson, E., & Sankar, J. S. (2012). Re-examining the Manning hypothesis:androgen receptor polymorphism and the 2D:4D digit ratio. Evolution and Human Behavior, 33, 557-561.

52.

Hankinson, S. E., Willett, W. C., Manson, J. E., Colditz, G. A., Hunter, D. J., Spiegelman, D., ... & Speizer, F. E. (1998). Plasma sex steroid hormone levels and risk of breast cancer in postmenopausal women. Journal of the National Cancer Institute, 90, 1292-1299.

53.

Harris, C. R., Jenkins, M., & Glaser, D. (2006). Gender differences in risk assessment: Why do women take fewer risks than men. Judgment and Decision Making, 1, 48-63.

54.

Helleday, J., Bartfai, A., Ritzen, M., & Forsman, M. (1994). General intelligence and cognitive profile in women with congenital adrenal hyperplasia (CAH). Psychoneuroendocrinology, 9, 343-356.

55.

Honekopp, J., Manning, J. T., & Muller, C. (2006). Digit ratio (2D:4D) and physical fitness in males and females: Evidence for effects of prenatal androgens on sexually selected traits. Hormones and Behavior, 49, 545-549.

56.

Honekopp, J., & Schuster, M. (2010). A meta-analysis on 2D:4D and athletic prowess:Substantial relationships but neither hand out-predicts the other. Personality and Individual Differences, 48, 4-10.

57.

Honekopp, J., & Watson, S. (2011). Meta-analysis of the relationship between digit-ratio 2D: 4D and aggression. Personality and Individual Differences, 51, 381-386.

58.

Hurd, P. L., Vaillancourt, K. L., & Dinsdale, N. L. (2011). Aggression, digit ratio and variation in androgen receptor and monoamine oxidase A genes in men. Behavior Genetics, 41, 543-556.

59.

Josephs, R. A., Sellers, J. G., Newman, M. L., & Mehta, P. H. (2006). The mismatch effect:when testosterone and status are at odds. Journal of Personality and Social Psychology, 90, 999-1013.

60.

Joyce, C. W., Kelly, J. C., Chan, J. C., Colgan, G., O’Briain, D., Mc Cabe, J. P., & Curtin, W. (2013). Second to fourth digit ratio confirms aggressive tendencies in patients with boxers fractures. Injury, 44, 1636-1639.

61.

Jung, H., Kim, K. H., Yoon, S. J., & Kim, T. B. (2011). Second to fourth digit ratio: a predictor of prostate‐specific antigen level and the presence of prostate cancer. BJU International, 107, 591-596.

62.

Kallai, J., Csatho, A., Kover, F., Makany, T., Nemes, J., Horvath, K., Kovacs, N., Manning, J. T., Nadel, L., & Nagy, F. (2005). MRI-assessed volume of left and right hippocampi in females correlates with the relative length of the second and fourth fingers (the 2D:4D ratio). Psychiatry Research, 140, 199-210.

63.

Kallman, D. A., Plato, C. C., & Tobin, J. D. (1990). The role of muscle loss in the age-related decline of grip strength: Crosssectional and longitudinal perspectives. Journal of Gerontology, 45, 82-88.

64.

Kasperk, C. H., Wakley, G. K., Hierl, T., & Ziegler, R. (1997). Gonadal and adrenal androgens are potent regulators of human bone cell metabolism in vitro. Journal of Bone and Mineral Research, 12, 464-471.

65.

Kelley, D. B. (1993). Androgens and brain development: Possible contributions to developmental dyslexia. In A. M. Galaburda (Ed.), Dyslexia and development: Neurobiological aspects of extra-ordinary brains (pp.21-41). Cambridge, MA: Harvard University Press.

66.

Kelso, W. M., Nicholls, M. E., Warne, G. L., & Zacharin, M. (2000). Cerebral lateralization and cognitive functioning in patients with congenital adrenal hyperplasia. Neuropsychology, 14, 370-378.

67.

Kempel, P., Gohlke, B., Klempau, J., Zinsberger, P., Reuter, M., & Hennig, J. (2005). Second to fourth digit length, testosterone, and spatial ability. Intelligence, 3, 215-230.

68.

Keshavan, M. S., & Hogarty, G. E. (1999). Brain maturational processes and delayed onset in schizophrenia. Development and Psychopathology, 11, 525-543.

69.

Klump, K. L., Gobrogge, K. L., Perkins, P. S., Thorne, D., Sisk, C. L., & Breedlove, S. (2006). Preliminary evidence that gonadal hormones organize and activate disordered eating. Psychological Medicine, 36, 539-546.

70.

Knickmeyer, R., Baron‐Cohen, S., Raggatt, P., & Taylor, K. (2005). Foetal testosterone, social relationships, and restricted interests in children. Journal of Child Psychology and Psychiatry, 46, 198-210.

71.

Kritz-Silverstein, D., & Barrett-Connor, E. (1994). Grip strength and bone mineral density in older women. Journal of Bone Mineral Research, 9, 45-51.

72.

Livshits, G., & Kobyliansky, E. (1991). Fluctuating asymmetry as a possible measure of developmental homeostasis in humans: A review. Human Biology, 63, 441-446.

73.

Logan, S., & Johnston, R. (2010). Investigating gender differences in reading. Educational Review, 62, 175-187.

74.

Lombardo, M. V., Ashwin, E., Auyeung, B., Chakrabarti, B., Lai, M. C., Taylor, K., ... & Baron-Cohen, S. (2012). Fetal programming effects of testosterone on the reward system and behavioral approach tendencies in humans. Biological Psychiatry, 72, 839-847.

75.

Lutchmaya, S., Baron-Cohen, S., & Raggatt, P. (2002). Foetal testosterone and eye contact in 12-month-old human infants. Infant Behavior and Development, 25, 327-335.

76.

Lutchmaya S, Baron-Cohen S, Raggatt P, Knickmeyer R, Manning, J. T. (2004). 2nd to 4th digit ratios, fetal testosterone and estradiol. Early Human Development, 77, 23-28.

77.

Manikkam, M., Crespi, E. J., Doop, D. D., Herkimer, C. Lee, J. S., Yu, S., … Padmanabhan, V. (2004). Fetal programming:prenatal testosterone excess leads to fetal growth retardation and postnatal catch-up growth in sheep. Endocrinology, 145, 790-798.

78.

Manning, J. T. (2002). Digit ratio: A pointer to fertility, behavior, and health. New Jersey:Rutgers University Press.

79.

Manning, J. T. (2011). Resolving the role of prenatal sex steroids in the development of digit ratio. Proceedings of the National Academy of Sciences, 108, 16143-16144.

80.

Manning, J. T., Barley, L., Walton, J., Lewis-Jones, D. I., Trivers, R. L., Singh, D., ... & Szwed, A. (2000). The 2nd: 4th digit ratio, sexual dimorphism, population differences, and reproductive success: evidence for sexually antagonistic genes?. Evolution and Human Behavior, 21, 163-183.

81.

Manning, J. T., Baron-Cohen, S., Wheelwright, S., & Sanders, G. (2001). The 2nd to 4th digit ratio and autism. Developmental Medicine and Child Neurology, 43, 160-164.

82.

Manning, J. T., & Bundred, P. E. (2000). The ratio of 2nd to 4th digit length: a new predictor of disease predisposition? Medical Hypotheses, 54, 855-857.

83.

Manning, J. T., & Bundred, P. E. (2001). The ratio of second to fourth digit length and age at first myocardial infarction in men: a link with testosterone? British Journal of Cardiology, 8, 720-723.

84.

Manning, J. T., Bundred, P. E., Newton, D. J., & Flanagan, B. F. (2003). The second to fourth digit ratio and variation in the androgen receptor gene. Evolution and Human Behavior, 24, 399-405.

85.

Manning, J. T., Churchill, A. J., & Peters, M. (2007). The effects of sex, ethnicity, and sexual orientation on self-measured digit ratio (2D:4D). Archives of Sexual Behavior, 36, 223-233.

86.

Manning, J. T., & Leinster, S. J. (2001). re: The ratio of 2nd to 4th digit length and age at presentation of breast cancer: a link with prenatal oestrogen?. The Breast, 10, 355-357.

87.

Manning, J. T., Morris, L., & Caswell, N. (2007). Endurance running and digit ratio (2D:4D):implications for fetal testosterone effects on running speed and vascular health. American Journal of Human Biology, 19, 416-421.

88.

Manning, J. T., & Peters, M. (2009). Digit ratio (2D: 4D) and hand preference for writing in the BBC Internet Study. Laterality, 14, 528-540.

89.

Manning, J. T., Scutt, D., Wilson, J., & Lewis-Jones, D. I. (1998). The ratio of 2nd to 4th digit length: a predictor of sperm numbers and concentrations of testosterone, luteinizing hormone and oestrogen. Human Reproduction, 13, 3000-3004.

90.

Manning, J. T., & Taylor, R. P. (2001). Second to fourth digit ratio and male ability in sport:implications for sexual selection in humans. Evolution and Human Behavior, 22, 61-69.

91.

Manning, J. T., Trivers, R. L., Thornhill, R., & Singh, D. (2000). The 2nd:4th digit ratio and asymmetry of hand performance in Jamaican children. Laterality, 5, 121-132.

92.

Martel, M. M., Gobrogge, K. L., Breedlove, S. M., & Nigg, J. T. (2008). Masculinized finger-length ratios of boys, but not girls, are associated with attention-deficit/hyperactivity disorder. Behavioral Neuroscience, 122, 273-281.

93.

Martin, S. M., Manning, J. T., & Dowrick, C. F. (1999). Fluctuating asymmetry, relative digit length, and depression in men. Evolution and Human Behavior, 20, 203-214.

94.

Mazur, A., & Booth, A. (1998). Testosterone and dominance in men. Behavioral and Brain Sciences, 21, 353-363.

95.

McFadden, D., Westhafer, J. G., Pasanen, E. G., Carlson, C. L., & Tucker, D. M. (2005). Physiological evidence of hypermasculinization in boys with the inattentive type of attentiondeficit/hyperactivity disorder (ADHD). Clinical Neuroscience Research, 5, 233-245.

96.

McIntyre, M. H., Ellison, P. T., Lieberman, D. E., Demerath, E., & Towne, B. (2005). The development of sex differences in digital formula from infancy in the Fels Longitudinal Study. Proceedings of the Royal Society B:Biological Sciences, 272, 1473-1479.

97.

Medland, S. E., Zayats, T., Glaser, B., Nyholt, D. R., Gordon, S. D., Wright, M. J., ... & Evans, D. M. (2010). A Variant in LIN28B Is Associated with 2D: 4D Finger-Length Ratio, a Putative Retrospective Biomarker of Prenatal Testosterone Exposure. The American Journal of Human Genetics, 86, 519-525.

98.

Millet, K., & Dewitte, S. (2007). Digit ratio (2D:4D) moderates the impact of an aggressive music video on aggression. Personality and Individual Differences, 43, 289-294.

99.

Milne, E., White, S., Campbell, R., Swettenham, J., Hansen, P., & Ramus, F. (2006). Motion and form coherence detection in autistic spectrum disorder: Relationship to motor control and 2: 4 digit ratio. Journal of Autism and Developmental Disorders, 36, 225-237.

100.

Moller, A. P., & Swaddle, J. P. (1997). Asymmetry, Developmental Stability, and Evolution. Oxford:Oxford University Press.

101.

Moore, T., Quinter, C., & Freeman, L. M. (2005). Lack of correlation between 2D: 4D ratio and assertiveness in college age women. Personality and Individual Differences, 39, 115-121.

102.

Moreira, B., Matsushita, R., & Da Silva, S. (2010). Risk seeking behavior of preschool children in a gambling task. Journal of Economic Psychology, 31, 794-801.

103.

Muller, D. C., Baglietto, L., Manning, J. T., McLean, C., Hopper, J. L., English, D. R., Giles, G. G., and Severi, G. (2012). Second to fourth digit ratio (2D:4D), breast cancer risk factors, and breast cancer risk: a prospective cohort study. British Journal of Cancer, 107, 1631-1636.

104.

Nelson, R. J. (3rd Ed.). (2005). An Introduction to Behavioral Endocrinology. Sinauer Associates, Inc.

105.

Nelson, E., & Shultz, S. (2010). Finger length ratios (2D: 4D) in anthropoids implicate reduced prenatal androgens in social bonding. American Journal of Physical Anthropology, 141, 395-405.

106.

Nicholls, M. E., Orr, C. A., Yates, M. J., & Loftus, A. M. (2008). A new means of measuring index/ring finger (2D:4D) ratio and its association with gender and hand preference. Laterality, 13, 71-91.

107.

Noipayak, P. (2009). The ratio of 2nd and 4th digit length in autistic children. Medical Journal of the Medical Association of Thailand, 92, 1040.

108.

Okten, A., Kalyoncu, M., & Yaris, N. (2002). The ratio of second- and fourth-digit lengths and congenital adrenal hyperplasia due to 21-hydroxylase deficiency. Early Human Development, 70, 47-54.

109.

Padmanabhan, V., Manikkam, M., Recabarren, S., & Foster, D. (2006). Prenatal testosterone excess programs reproductive and metabolic dysfunction in the females. Molecular and Cellular Endocrinology, 246, 165-174.

110.

Palmer, A. R., & Strobeck, C. (1986). Fluctuating asymmetry: measurement, analysis and patterns. Annual Review of Ecology and Systematics, 17, 391-421.

111.

Page, S. T., Amory, J. K., Bowman, F. D., Anawalt, B. D., Matsumoto, A. M., Bremner, W. J., & Tenover, J. L. (2005). Exogenous testosterone (T) alone or with finasteride increases physical performance, grip strength, and lean body mass in older men with low serum T. Journal of Clinical Endocrinology &Metabolism, 90, 1502-1510.

112.

Parsons, P. A. (1990). Fluctuating asymmetry: an epigenetic measure of stress. Biological Reviews, 65, 131-145.

113.

Peters, M., Mackenzie, K., & Bryden, P. (2002). Finger length and distal finger extent patterns in humans. American Journal of Physical Anthropology, 117, 209-217.

114.

Putz, D. A., Gaulin, S. J. C., Sporter, R. J., & McBurney, D. H. (2004). Sex hormones and finger length: what does 2D:4D indicate? Evolution and Human Behavior, 25, 182-199.

115.

Puts, D. A., McDaniel, M. A., Jordan, C. L., & Breedlove, S. M. (2008). Spatial ability and prenatal androgens: Meta-analyses of congenital adrenal hyperplasia and digit ratio (2D:4D)studies. Archives of Sexual Behavior, 37, 100-111.

116.

Rahman, A. A., Lophatananon, A., Stewart-Brown, S., Harriss, D., Anderson, J., Parker, T., ... & Muir, K. (2010). Hand pattern indicates prostate cancer risk. British Journal of Cancer, 104, 175-177.

117.

Raymond, M., Pontier, D., Dufour, A. B., & Moller, A. P. (1996). Frequency-dependent maintenance of left handedness in humans. Proceedings of the Royal Society of London. Series B: Biological Sciences, 263, 1627-1633.

118.

Ronalds, G., Phillips, D. I. W., Godfrey, K. M., & Manning, J. T. (2002). The ratio of second to forth digit lengths: a marker of impaired fetal growth? Early Human Development, 68, 21-26.

119.

Roney, J. R., Whitham, J. C., Leoni, M., Bellem, A., Wielebnowski, N., & Maestripieri, D. (2004). Relative digit lengths and testosterone levels in Guinea baboons. Hormones and Behavior, 45, 285-290.

120.

Sapienza, P., Zingales, L., & Maestripieri, D. (2009). Gender differences in financial risk aversion and career choices are affected by testosterone. Proceedings of the National Academy of Sciences, 106, 15268-15273.

121.

Senju, A., & Johnson, M. H. (2009). The eye contact effect: mechanisms and development. Trends in Cognitive Sciences, 13, 127-134.

122.

Shoup, M. L., & Gallup, G. G. (2008). Men’s faces convey information about their bodies and their behavior: What you see is what you get. Evolutionary Psychology, 6, 469-479.

123.

Sim, K. (2013). The relationship between sextypical body shape and quality indicators. Journal of Social, Evolutionary, and Cultural Psychology, 7, 97-120.

124.

Singh, D. (1993). Adaptive significance of female physical attractiveness: Role of waist-to-hip ratio. Journal of Personality & Social Psychology, 65, 293-307.

125.

Smith, A. R., Hawkeswood, S. E., & Joiner, T. E. (2010). The measure of a man: associations between digit ratio and disordered eating in males. International Journal of Eating Disorders, 43, 543-548.

126.

Stenstrom, E., Saad, G., Nepomuceno, M. V., & Mendenhall, Z. (2011). Testosterone and domain-specific risk: Digit ratios (2D:4D and rel2) as predictors of recreational, financial, and social risk-taking behaviors. Personality and Individual Differences, 51, 412-416.

127.

Stevenson, J. C., Everson, P. M., Williams, D. C., Hipskind, G., Grimes, M., & Mahoney, E. R. (2007). Attention deficit/hyperactivity disorder (ADHD) symptoms and digit ratios in a college sample. American Journal of Human Biology, 19, 41-50.

128.

Talarovičová, A., Kršková, L., & Blažeková, J. (2009). Testosterone enhancement during pregnancy influences the 2D: 4D ratio and open field motor activity of rat siblings in adulthood. Hormones and Behavior, 55, 235-239.

129.

Tester, N., & Campbell, A. (2007). Sporting achievement: What is the contribution of digit ratio? Journal of Personality, 75, 663-678.

130.

Thornhill, R., & Gangestad, S. (1993). Human facial beauty: averageness, symmetry, and parasite resistance. Human Nature, 4, 237-270.

131.

Thornhill, R., & Gangestad, S. W. (1994). Human fluctuating asymmetry and sexual behavior. Psychological Science, 5, 297-302.

132.

Trichopoulos, D. (1990). Hypothesis: does breast cancer originate in utero?. The Lancet, 335, 939-940.

133.

van Anders, S. M., Vernon, P. A., & Wilbur, C. J. (2006). Finger-length ratios show evidence of prenatal hormone-transfer between oppositesex twins. Hormones and Behavior, 49, 315-319.

134.

van Valen, L. (1962). A study of fluctuating asymmetry. Evolution, 16, 125-142.

135.

Vermeersch, H., T'Sjoen, G., Kaufman, J. M., & Vincke, J. (2008). 2d: 4d, sex steroid hormones and human psychological sex differences. Hormones and Behavior, 54, 340-346.

136.

Voracek, M., & Dressler, S. G. (2006). Lack of correlation between digit ratio (2D: 4D) and Baron-Cohen’s “Reading the Mind in the Eyes” test, empathy, systemising, and autism-spectrum quotients in a general population sample. Personality and Individual Differences, 41, 1481-1491.

137.

Voracek, M., & Loibl, L. M. (2009). Scientometric analysis and bibliography of digit ratio (2D:4D) research, 1998-2008 1, 2. Psychological Reports, 104, 922-956.

138.

Voracek, M., Manning, J. T., & Dressler, S. G. (2007). Repeatability and interobserver error of digit ratio (2D: 4D) measurements made by experts. American Journal of Human Biology, 19, 142-146.

139.

Voracek, M., Reimer, B., Ertl, C., & Dressler, S. G. (2006). Digit ratio (2D:4D), lateral preferences, and performance in fencing 1. Perceptual and Motor Skills, 103, 427-446.

140.

Voyer, D., Voyer, S., & Bryden, M. P. (1995). Magnitude of sex differences in spatial abilities: a meta-analysis and consideration of critical variables. Psychological Bulletin, 117, 250-270.

141.

Weber, D., Skirbekk, V., Freund, I., & Herlitz, A. (2014). The changing face of cognitive gender differences in Europe. Proceedings of the National Academy of Sciences, 111, 11673-11678.

142.

Weissman, M. M., & Klerman, G. L. (1977). Sex differences and the epidemiology of depression. Archives of General Psychiatry, 34, 98-11.

143.

Wisniewski, A. B. (1998). Sexually-dimorphic patterns of cortical asymmetry, and the role for sex steroid hormones in determining cortical patterns of lateralization. Psychoneuroendocrinology, 23, 519-547.

144.

Zheng, Z., & Cohn, M. J. (2011). Developmental basis of sexually dimorphic digit ratios. Proceedings of the National Academy of Sciences, 108, 16289-16294.

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