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生態學 1926

捕食者與獵物:洛特卡—沃爾泰拉方程

阿爾弗雷德·洛特卡 與 維托·沃爾泰拉

兩個耦合方程,讓捕食者與獵物的數量永遠此消彼長,一個峰追著另一個峰。

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In depth · the introduction

一片沒有狐狸的草地會擠滿兔子;一片狐狸太多的草地則會兔狐俱空。洛特卡與沃爾泰拉找到了這場永不停歇的蹺蹺板背後那條簡單的規則。

把這個想法拆開看

想象兩個彼此推搡的數。獵物多時,捕食者吃得好、繁殖快。可捕食者一多,被吃的獵物也多,於是獵物崩落。沒什麼可吃,捕食者隨之挨餓、減少——而捕食者一少,獵物又反彈,整個循環重新開始。

洛特卡與沃爾泰拉用兩個簡短的方程恰好抓住了這一切:獵物自行增長,遇到捕食者則減少;捕食者遇到獵物則增長,自行則減少。讓方程跑起來,兩個種群永遠安定不下來——它們永遠以波浪起落,捕食者的波總比獵物的波慢半拍。這是頭一回,有人把食物鏈裡的你來我往,化成了真能做出預測的數學。

一個魚市謎題

把這個問題交到意大利數學家維托·沃爾泰拉手上的,是他的女婿、研究亞得里亞海漁獲的生物學家翁貝托·丹科納。丹科納在市場記錄裡注意到一樁怪事:第一次世界大戰期間,漁船都停在港裡,捕上來的魚反而更多是鯊與鰩——那些捕食者——而不是你以為會恢復的食用魚。為什麼捕得少,反倒利於獵手?沃爾泰拉造出他的方程來尋找答案,方程給出一個乾淨的回答:對捕食者與獵物一視同仁的捕撈,其實是在撐著捕食者;一旦停捕,平衡便朝它們那邊倒回去。沃爾泰拉並不知道,美國的化學家兼精算師阿爾弗雷德·洛特卡幾年前已寫下同樣的方程,他是借振盪的化學反應作類比推出來的。兩人通信、釐清了先後,此後這個模型便一直冠以二人之名。

它為何重要

在此之前,生態學是一門靠細緻觀察與文字的科學——達爾文的「生存鬥爭」生動,卻無法計算。洛特卡與沃爾泰拉表明,物種之間那糾纏的你來我往,可以服從精確到足以預測、甚至足以出人意料的方程:捕魚竟有利於捕食者,這絕非常識會遞給你的結論。僅這一個示範,就打開了為整個生態系統、疾病傳播、漁業與蟲害治理建模的大門。

一架總在遲到的蹺蹺板

設想一架其中一頭無法即時反應的蹺蹺板。把「獵物」那頭壓下去,「捕食者」那頭要過一陣才升起;等捕食者升到高處,獵物其實已經在下沉了;然後捕食者也下沉——但遲了,總是遲一步。正是這內建的延遲,讓兩頭不停振盪,而非歸於平衡。捕食者的峰值永遠在環上追著獵物的峰值,慢四分之一圈,卻永遠追不上。

兩條隨時間振盪的曲線,綠色獵物與赭色捕食者,捕食者的峰值滯後於獵物的峰值;捕撈滑桿抬高時,獵物的平均水平上升、捕食者的下降。

之前與之後

馬爾薩斯(1798)已表明,一個種群若放任不管便會爆炸式增長;韋爾許爾斯特用一個上限把這增長壓彎。洛特卡與沃爾泰拉補上了缺失的一塊——當兩個這樣的種群相互取食時會發生什麼——於是給達爾文(1859)只用文字敘述過的那場鬥爭配上了方程。他們的循環成了疾病模型與漁業科學的祖先;而他們那脆弱、完美平衡的環——任何一點添加的現實都能輕易擾亂——預示了洛倫茨(1963)後來在混沌理論裡使之聞名的、對微小變化的敏感。

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