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第三章 HPG 轴、睾酮与挑战假说

从鸟类开始

下丘脑释放促性腺激素释放激素(GnRH),垂体释放黄体生成素(LH),LH 促使睾丸(女性主要是卵巢和肾上腺)合成睾酮。睾酮反过来抑制 GnRH 和 LH,这是另一条负反馈回路:下丘脑—垂体—性腺轴(HPG 轴)。

证据

温菲尔德(Wingfield)、赫格纳(Hegner)、达夫蒂(Dufty)与鲍尔(Ball)在 1990 年提出「挑战假说」:在一夫一妻、雄鸟参与育雏的鸟类中,繁殖季节的睾酮通常维持在足以支持繁殖的较低基线;只有在面临领地入侵、争夺配偶等社会挑战时,睾酮才升高到更高水平,并在挑战过去后回落。持续的高睾酮代价高昂,会干扰育雏 (Wingfield et al. 1990)。

这个假说的要点不是「睾酮越高越好」,而恰恰相反:睾酮是一个随情境上下调节的信号,高水平只在需要时出现。阿彻(Archer)把它用于人类,发现人类数据部分符合:睾酮在竞争前后会变化,基线睾酮与攻击性的相关很弱 (Archer 2006)。

人类证据有多大

关于睾酮,流行说法与实证效应量之间差距很大。

证据

艾森内格(Eisenegger)等人在双盲安慰剂对照的最后通牒博弈中给女性受试者服用睾酮。服用睾酮者提出的分配更公平,而不是更自私;但相信自己服用了睾酮的人(无论实际服用的是什么)提出的分配更不公平。作者的解释是:睾酮可能促进追求地位的行为,而在这种情境中,公平的提议有助于维护地位;民间关于睾酮使人自私好斗的信念本身会改变行为 (Eisenegger et al. 2010)。艾森内格、豪斯霍费尔(Haushofer)与费尔(Fehr)随后在综述中提出,睾酮的作用应理解为促进地位寻求,而不是单纯促进攻击 (Eisenegger et al. 2011)。

证据

关于「赢家睾酮上升、输家下降」的「胜负效应」,赫尼奥莱(Geniole)等人的更新元分析发现,赢家在比赛后的睾酮确实略高于输家,但效应小,在实验室比赛中尤其弱,在真实体育比赛中较大,各研究之间差异很大 (Geniole et al. 2017)。

证据

「力量姿势」的流行说法认为,摆两分钟扩张性的姿势就能升高睾酮、降低皮质醇。拉内希尔(Ranehill)等人在 200 人的较大样本中重复这一实验,没有发现姿势对睾酮或皮质醇的影响 (Ranehill et al. 2015)。

为了直观地理解「效应小」意味着什么,可以把标准化效应量换算成两个更具体的数字。

推导

效应量的直观含义:两组之间标准化差异为 dd 时,从高的一组随机抽一人、从低的一组随机抽一人,前者确实更高的概率是 Φ(d/2)\Phi(d/\sqrt{2});两个分布的重叠比例是 2Φ(−|d|/2)2\Phi(-|d|/2)。

# What small hormone-behaviour effects mean in practice.
import math
from statistics import NormalDist
N = NormalDist()

def from_d(d):
    """Standardised mean difference -> chance that a random member of group 1 scores higher than a
    random member of group 2 (common-language effect size), and the overlap of the two distributions."""
    cl = N.cdf(d / 2 ** 0.5)
    overlap = 2 * N.cdf(-abs(d) / 2)
    return cl, overlap

print("d      P(random member of higher group is higher)   overlap of distributions")
for d in (0.1, 0.2, 0.5, 0.8):
    cl, ov = from_d(d)
    print(f"{d:4.1f}   {100 * cl:5.1f} %                                     {100 * ov:5.1f} %")

r = -0.061          # testosterone x cortisol interaction, Dekkers et al. 2019 meta-analysis
print(f"\nr = {r}: explains {100 * r * r:.2f} % of the variance in status-relevant behaviour")
# People needed to detect such an effect once, two-sided alpha 0.05, 80 % power (Fisher z approximation)
z = 0.5 * math.log((1 + abs(r)) / (1 - abs(r)))
n = ((N.inv_cdf(0.975) + N.inv_cdf(0.80)) / z) ** 2 + 3
print(f"participants needed for 80 % power to detect r = {abs(r)}: {math.ceil(n)}")
# d      P(random member of higher group is higher)   overlap of distributions
#  0.1    52.8 %                                      96.0 %
#  0.2    55.6 %                                      92.0 %
#  0.5    63.8 %                                      80.3 %
#  0.8    71.4 %                                      68.9 %
#
# r = -0.061: explains 0.37 % of the variance in status-relevant behaviour
# participants needed for 80 % power to detect r = 0.061: 2108

皮质醇抑制睾酮

HPA 轴与 HPG 轴并不独立。

证据

维奥(Viau)综述了两条轴之间的双向联系:在动物中,应激和糖皮质激素可在下丘脑、垂体和睾丸多个层面抑制生殖轴;而性激素也调节 HPA 轴对应激的反应 (Viau 2002)。

梅塔(Mehta)与约瑟夫斯(Josephs)由此提出「双激素假说」:睾酮与支配、地位行为的关联,只在皮质醇低的人身上出现;皮质醇高时,这种关联消失或反转 (Mehta and Josephs 2010)。

证据

德克尔斯(Dekkers)等人对双激素假说做了元分析,发现睾酮与皮质醇的交互作用对地位相关行为虽在统计上显著,但非常小(r=−0.061r = -0.061),并且存在发表偏倚、分析灵活性和单个研究检验力不足的问题。作者认为该假说只得到「边缘」支持,需要大型预注册研究 (Dekkers et al. 2019)。

上面的程序已经算出:r=−0.061r = -0.061 只解释约 0.4% 的变异,要以 80% 的检验力检出它一次,需要约两千一百人。大多数单个研究的样本远小于此。

挑战评估与威胁评估

同一个压力情境,可以被评估为挑战,也可以被评估为威胁。

证据

布拉斯科维奇(Blascovich)与门德斯(Mendes)的生物心理社会模型认为,当人评估自己的资源足以应对任务要求时,处于「挑战」状态:心输出量上升、外周血管阻力下降;当评估要求超过资源时,处于「威胁」状态:心输出量变化较小、血管阻力上升 (Blascovich and Mendes 2000)。贾米森(Jamieson)等人发现,在考试或社会压力测试前,引导受试者把身体的唤醒重新理解为「有助于表现的」,能使心血管反应更接近挑战模式 (Jamieson et al. 2012)。

推导

一个耦合两条轴的玩具模型:60 分钟的比赛中,皮质醇随评估到的要求上升;睾酮的合成在「可以赢的挑战」中得到增强,并受高皮质醇抑制。所有量为任意单位(静息为 1),参数纯属示意。

# Toy model of two coupled axes during a 60-minute contest (arbitrary units, rest = 1).
# Cortisol C rises with appraised demand; testosterone T production is boosted by a contest the person
# appraises as a winnable challenge and is inhibited by high cortisol (HPA -> HPG cross-talk, Viau 2002).
# Parameters are illustrative; effect sizes in real people are much smaller and more variable (see text).
def contest(c_drive, t_boost, minutes=240, dt=0.5):
    C = T = 1.0
    hC, hT, KI = 1 / 60, 1 / 45, 2.0          # clearance per minute; inhibition constant
    out, t = {}, 0.0
    while t <= minutes:
        on = t < 60
        dC = hC * ((1 + (c_drive if on else 0)) - C)
        # production: boosted during a challenge, inhibited by cortisol (equals 1 at C = 1 without a boost)
        prod = (1 + (t_boost if on else 0)) * (1 + (1 / KI) ** 2) / (1 + (C / KI) ** 2)
        dT = hT * (prod - T)
        C += dt * dC; T += dt * dT
        for m in (30, 60, 120, 240):
            if abs(t - m) < dt / 2:
                out[m] = (C, T)
        t += dt
    return out

print(f"{'appraisal':32s}  min     C     T   T/C")
for name, cd, tb in (("challenge (demands <= resources)", 0.8, 0.4),
                     ("threat (demands > resources)", 2.5, 0.0),
                     ("no contest", 0.0, 0.0)):
    o = contest(cd, tb)
    for m in (60, 120, 240):
        C, T = o[m]
        print(f"{name if m == 60 else '':32s} {m:4d}: {C:5.2f} {T:5.2f} {T / C:5.2f}")
# appraisal                         min     C     T   T/C
# challenge (demands <= resources)   60:  1.50  1.15  0.76
#                                   120:  1.18  0.95  0.81
#                                   240:  1.02  0.98  0.95
# threat (demands > resources)       60:  2.57  0.71  0.28
#                                   120:  1.58  0.68  0.43
#                                   240:  1.08  0.92  0.85
# no contest                         60:  1.00  1.00  1.00
#                                   120:  1.00  1.00  1.00
#                                   240:  1.00  1.00  1.00

在模型里,挑战评估下睾酮小幅上升、皮质醇中度上升,两小时内恢复;威胁评估下皮质醇大幅上升,睾酮被压低,并在比赛结束后仍低一段时间。这是对维奥所述交叉抑制与布拉斯科维奇所述两种评估的简化模型,不是对人体激素量的预测:上面引用的人类研究显示,真实效应远小于、也远比模型中的曲线多变。

注

本书不讨论、也不建议任何睾酮或其他激素的补充、处方外使用或「天然提升」方法。睾酮治疗是有适应证和风险的医疗措施,只能由医生决定。

假说 · 尚未验证

尚未确立。 在准备一次重要的公开报告时,接受五分钟「唤醒重评」指导(把心跳加快理解为身体在为表现做准备)的人,比接受同等时长中性指导的人,报告前后唾液睾酮与皮质醇之比下降得更少。如果在预注册的随机试验中,两组比值变化之差的 95% 置信区间完全落在预先设定的等效界限 ±0.1(对数单位)之内,假说就被推翻。

小结

睾酮是随社会情境上下调节的信号,不是「自信激素」。人类研究中,它对行为的效应小、依赖情境,常被流行说法夸大。皮质醇与睾酮之间存在生理上的交叉抑制,但双激素假说的人类证据目前很弱。比激素数值本身更可以由人掌握的,是对情境的评估。

参考文献

Archer, John. 2006. “Testosterone and Human Aggression: An Evaluation of the Challenge Hypothesis.” Neuroscience and Biobehavioral Reviews 30 (3): 319–45.
Blascovich, Jim, and Wendy Berry Mendes. 2000. “Challenge and Threat Appraisals: The Role of Affective Cues.” In Feeling and Thinking: The Role of Affect in Social Cognition, edited by Joseph P. Forgas. Cambridge University Press.
Dekkers, Tycho J., Joost A. Agelink van Rentergem, Bart Meijer, Arne Popma, Eline Wagemaker, and Hilde M. Huizenga. 2019. “A Meta-Analytical Evaluation of the Dual-Hormone Hypothesis: Does Cortisol Moderate the Relationship Between Testosterone and Status, Dominance, Risk Taking, Aggression, and Psychopathy?” Neuroscience and Biobehavioral Reviews 96: 250–71.
Eisenegger, Christoph, Johannes Haushofer, and Ernst Fehr. 2011. “The Role of Testosterone in Social Interaction.” Trends in Cognitive Sciences 15 (6): 263–71.
Eisenegger, Christoph, Michael Naef, Romana Snozzi, Markus Heinrichs, and Ernst Fehr. 2010. “Prejudice and Truth about the Effect of Testosterone on Human Bargaining Behaviour.” Nature 463 (7279): 356–59.
Geniole, Shawn N., Brian M. Bird, Erika L. Ruddick, and Justin M. Carré. 2017. “Effects of Competition Outcome on Testosterone Concentrations in Humans: An Updated Meta-Analysis.” Hormones and Behavior 92: 37–50.
Jamieson, Jeremy P., Matthew K. Nock, and Wendy Berry Mendes. 2012. “Mind over Matter: Reappraising Arousal Improves Cardiovascular and Cognitive Responses to Stress.” Journal of Experimental Psychology: General 141 (3): 417–22.
Mehta, Pranjal H., and Robert A. Josephs. 2010. “Testosterone and Cortisol Jointly Regulate Dominance: Evidence for a Dual-Hormone Hypothesis.” Hormones and Behavior 58 (5): 898–906.
Ranehill, Eva, Anna Dreber, Magnus Johannesson, Susanne Leiberg, Sunhae Sul, and Roberto A. Weber. 2015. “Assessing the Robustness of Power Posing: No Effect on Hormones and Risk Tolerance in a Large Sample of Men and Women.” Psychological Science 26 (5): 653–56.
Viau, Victor. 2002. “Functional Cross-Talk Between the Hypothalamic-Pituitary-Gonadal and -Adrenal Axes.” Journal of Neuroendocrinology 14 (6): 506–13.
Wingfield, John C., Robert E. Hegner, Alfred M. Dufty, and Gregory F. Ball. 1990. “The ‘Challenge Hypothesis’: Theoretical Implications for Patterns of Testosterone Secretion, Mating Systems, and Breeding Strategies.” The American Naturalist 136 (6): 829–46.

Fan, Y. (2026). Dynamical Systems of Neurotransmitters and Endocrinology: Dopamine Reward Prediction Error, Serotonergic Homeostasis, HPA-Axis Allostatic Load and the Challenge Hypothesis, Read Beside East Asian Classics of Mind Regulation [神经递质与内分泌的动力系统] (Version 1.0.0). 5D Library. https://doi.org/10.5281/zenodo.23738782 · Licensed under Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International (CC BY-NC-ND 4.0).