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生态学 1926

捕食者与猎物:洛特卡—沃尔泰拉方程

阿尔弗雷德·洛特卡 与 维托·沃尔泰拉

两个耦合方程,让捕食者与猎物的数量永远此消彼长,一个峰追着另一个峰。

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In depth · the introduction

一片没有狐狸的草地会挤满兔子;一片狐狸太多的草地则会兔狐俱空。洛特卡与沃尔泰拉找到了这场永不停歇的跷跷板背后那条简单的规则。

把这个想法拆开看

想象两个彼此推搡的数。猎物多时,捕食者吃得好、繁殖快。可捕食者一多,被吃的猎物也多,于是猎物崩落。没什么可吃,捕食者随之挨饿、减少——而捕食者一少,猎物又反弹,整个循环重新开始。

洛特卡与沃尔泰拉用两个简短的方程恰好抓住了这一切:猎物自行增长,遇到捕食者则减少;捕食者遇到猎物则增长,自行则减少。让方程跑起来,两个种群永远安定不下来——它们永远以波浪起落,捕食者的波总比猎物的波慢半拍。这是头一回,有人把食物链里的你来我往,化成了真能做出预测的数学。

一个鱼市谜题

把这个问题交到意大利数学家维托·沃尔泰拉手上的,是他的女婿、研究亚得里亚海渔获的生物学家翁贝托·丹科纳。丹科纳在市场记录里注意到一桩怪事:第一次世界大战期间,渔船都停在港里,捕上来的鱼反而更多是鲨与鳐——那些捕食者——而不是你以为会恢复的食用鱼。为什么捕得少,反倒利于猎手?沃尔泰拉造出他的方程来寻找答案,方程给出一个干净的回答:对捕食者与猎物一视同仁的捕捞,其实是在撑着捕食者;一旦停捕,平衡便朝它们那边倒回去。沃尔泰拉并不知道,美国的化学家兼精算师阿尔弗雷德·洛特卡几年前已写下同样的方程,他是借振荡的化学反应作类比推出来的。两人通信、厘清了先后,此后这个模型便一直冠以二人之名。

它为何重要

在此之前,生态学是一门靠细致观察与文字的科学——达尔文的「生存斗争」生动,却无法计算。洛特卡与沃尔泰拉表明,物种之间那纠缠的你来我往,可以服从精确到足以预测、甚至足以出人意料的方程:捕鱼竟有利于捕食者,这绝非常识会递给你的结论。仅这一个示范,就打开了为整个生态系统、疾病传播、渔业与虫害治理建模的大门。

一架总在迟到的跷跷板

设想一架其中一头无法即时反应的跷跷板。把「猎物」那头压下去,「捕食者」那头要过一阵才升起;等捕食者升到高处,猎物其实已经在下沉了;然后捕食者也下沉——但迟了,总是迟一步。正是这内建的延迟,让两头不停振荡,而非归于平衡。捕食者的峰值永远在环上追着猎物的峰值,慢四分之一圈,却永远追不上。

两条随时间振荡的曲线,绿色猎物与赭色捕食者,捕食者的峰值滞后于猎物的峰值;捕捞滑块抬高时,猎物的平均水平上升、捕食者的下降。

之前与之后

马尔萨斯(1798)已表明,一个种群若放任不管便会爆炸式增长;韦尔许尔斯特用一个上限把这增长压弯。洛特卡与沃尔泰拉补上了缺失的一块——当两个这样的种群相互取食时会发生什么——于是给达尔文(1859)只用文字叙述过的那场斗争配上了方程。他们的循环成了疾病模型与渔业科学的祖先;而他们那脆弱、完美平衡的环——任何一点添加的现实都能轻易扰乱——预示了洛伦茨(1963)后来在混沌理论里使之闻名的、对微小变化的敏感。

The original document
Original source text
Vito Volterra · Nature 118: 558–560 · 1926
The question (paraphrase)
Volterra's son-in-law, the marine biologist Umberto d'Ancona, had compiled fish-market statistics from the upper Adriatic across the years around the First World War. He noticed something odd: during and just after the war, when fishing had nearly stopped, the proportion of predatory fish (selachians — sharks and rays) in the catch rose, while food fish fell back. Less fishing favoured the predators. He asked Volterra whether mathematics could explain it.
The model (paraphrase)
Volterra treated the prey population x and the predator population y as continuous quantities. Left alone, prey would multiply without limit (rate α); predators, with nothing to eat, would die out (rate γ). The two are coupled only through encounters, taken proportional to the product x·y: such meetings remove prey (rate β) and feed predators (rate δ). This gives two coupled rate equations:
dx/dt = α·x − β·x·y and dy/dt = δ·x·y − γ·y. They have no fixed resting state except the balance point (x*, y*) = (γ/δ, α/β); away from it the two numbers circle that point forever, the predators peaking a quarter-cycle behind the prey.
The three laws (paraphrase)
Law of the periodic cycle — the two populations fluctuate periodically, the period set by the coefficients and the starting amounts. Law of conservation of the averages — averaged over a full cycle, each population sits exactly at its balance point, no matter how wild the swings. Law of the disturbance of the averages — if both species are destroyed in proportion to their numbers (uniform fishing), the average of the prey goes up and the average of the predator goes down. That last law answered d'Ancona: cutting the fishing of wartime had pushed the balance the other way, toward the predators.
[ … ]
Vito Volterra · Rome · 1926