And in particular, how the simple parts of organizations

我研究螞蟻，因為我喜歡思考組織的運作方式，

interact to create the behavior of the whole organization.

尤其想知道，單一的個體是如何透過互動的方式，

So, ant colonies are a good example of an organization like that,

就能創造出複雜完整的組織行為。

and there are many others. The web is one.

而蟻群就是最好的例子，

There are many biological systems like that --

其它像是網路，

brains, cells, developing embryos.

很多的生物系統，像是

There are about 10,000 species of ants.

大腦、細胞、發育中的胚胎都是類似的例子。

They all live in colonies consisting of one or a few queens,

螞蟻大約有一萬多種品種，

and then all the ants you see walking around are sterile female workers.

他們的蟻群都會有一隻或幾隻蟻后，

And all ant colonies have in common that there's no central control.

而那些走來走去的螞蟻，都是沒有生育能力的雌性工蟻，

Nobody tells anybody what to do.

而且所有蟻群都有一個共同點，就是他們沒有中央領導。

The queen just lays the eggs. There's no management.

沒有任何一隻工蟻被告知該做什麼事，

No ant directs the behavior of any other ant.

蟻后只顧生蛋，她並不做任何管理的事情，

And I try to figure out how that works.

沒有任何一隻螞蟻指揮其它螞蟻該做什麼事，

And I've been working for the past 20 years

我想試著了解他們是怎麼運作的，

on a population of seed-eating ants in southeastern Arizona.

所以我過去二十年，

Here's my study site. This is really a picture of ants,

一直在亞利桑那州的東南部，研究一種會吃種子的螞蟻，

and the rabbit just happens to be there.

這就是我研究的地方。這真的是一張螞蟻的照片，

And these ants are called harvester ants because they eat seeds.

只是剛好有隻兔子在那。

This is the nest of the mature colony, and there's the nest entrance.

這些螞蟻被稱作收割蟻，因為他們吃種子。

And they forage maybe for about 20 meters away,

這是一個發展健全的蟻巢，而且那裡有一個入口，

gather up the seeds and bring them back to the nest, and store them.

裡面的螞蟻們可能會到20公尺遠的地方覓食，

And every year I go there and make a map of my study site.

蒐集種子並且把它們帶回巢穴儲存。

This is just a road. And it's not very big:

每一年我都會去那裡，製作一張蟻群分佈的地圖，

it's about 250 meters on one side, 400 on the other.

只是一條路的範圍，不是很大，

And every colony has a name, which is a number,

大約有250公尺長，400公尺寬。

which is painted on a rock. And I go there every year

這裡的每個蟻群都有各自的編號，

and look for all the colonies that were alive the year before,

編號被標記在石頭上，所以每年我都可以回去那裡，

and figure out which ones have died, and put all the new ones on the map.

找出所有蟻群過去一年居住的位置，

And by doing this I know how old they all are.

然後觀察哪些蟻群已經滅絕，並在地圖上標註出新蟻群。

And because of that, I've been able to study how their behavior changes

持續透過這樣的工作，我可以知道這些蟻群活了多久，

as the colony gets older and larger.

也因為如此，隨著蟻群的成長及擴張，

So I want to tell you about the life cycle of a colony.

我可以研究他們的行為變化。

Ants never make more ants; colonies make more colonies.

我想先來談談關於蟻群的生命週期。

And they do that by each year sending out the reproductives --

絕大部分的螞蟻不能繁殖，但蟻群卻可以生產更多蟻群，

those are the ones with wings -- on a mating flight.

這是因為他們每年都會派出那些有翅膀的生殖蟻，

So every year, on the same day -- and it's a mystery exactly how that happens --

飛到蟻穴之外交配。

each colony sends out its virgin, unmated queens with wings, and the males,

而且很神奇的，都剛好在每年的同一天，

and they all fly to a common place. And they mate.

每個蟻群都會在那天送出長有翅膀的新蟻后與公蟻，

And this shows a recently virgin queen. Here's her wings.

一起飛到同個地方交配。

And she's in the process of mating with this male,

這是一隻最近拍攝到的新蟻后，那是她的翅膀，

and there's another male on top waiting his turn.

她正在跟這隻公蟻交配，

Often the queens mate more than once.

而且還有另一隻公蟻在上面等，等待輪到他交配。

And after that, the males all die. That's it for them.

通常每隻新蟻后都會交配一次以上，

(Laughter)

交配完，公蟻就會死，那就是他們的用處。

And then the newly mated queens fly off somewhere, drop their wings,

(笑聲)

dig a hole and go into that hole and start laying eggs.

接著這些懷孕的新蟻后會飛到某個地方，褪去她們的翅膀，

And they will live for 15 or 20 years, continuing to lay eggs

挖一個洞，鑽進去，然後開始產卵。

using the sperm from that original mating.

她們會活15到20年，並且利用與公蟻交配時留下的精液，

So the queen goes down in there.

每年持續不斷地產卵。

She lays eggs, she feeds the larvae -- so an ant starts as an egg, then it's a larva.

女王由此誕生。

She feeds the larvae by regurgitating from her fat reserves.

螞蟻是卵生的，女王會持續地產卵並扶養幼蟲，

Then, as soon as the ants -- the first group of ants -- emerge,

她利用反芻的方式，將體內豐富的貯存物餵食給幼蟲，

they're larvae. Then they're pupae. Then they come out as adult ants.

然後，她的第一批子民開始湧現，

They go out, they get the food, they dig the nest,

他們很快地從幼蟲變成蛹，然後破繭而出變為成蟻。

and the queen never comes out again.

這些成蟻會出去找食物，或蓋巢穴，

So this is a one-year-old colony -- this happens to be 536.

而女王從此不再踏出她的王宮。

There's the nest entrance, there's a pencil for scale.

這是一個一歲大的蟻群，它的編號是536，

So this is the colony founded by a queen the previous summer.

那裡有一個巢穴的入口，鉛筆是比例尺，

This is a three-year-old colony.

這是由某隻蟻后在去年夏天所建立的。

There's the nest entrance, there's a pencil for scale.

這是一個三歲大的蟻群，

They make a midden, a pile of refuse -- mostly the husks of the seeds that they eat.

巢穴的入口在那裡，鉛筆是比例尺，

This is a five-year-old colony. This is the nest entrance, here's a pencil for scale.

他們在那裡丟了一堆廢棄物，大部分是吃剩的穀殼。

This is about as big as they get, about a meter across.

這是一個五歲大的蟻群，那是巢穴入口，鉛筆是比例尺，

And then this is how colony size and numbers of worker ants changes --

這蟻群已成長至最大，大約一公尺長。

so this is about 10,000 worker ants --

這張圖是用來觀察蟻群大小和工蟻數的變化，

changes as a function of colony age, in years.

所以這(五歲大的)蟻群大約有一萬隻工蟻，

So it starts out with zero ants, just the founding queen,

蟻群大小的變化是蟻群年紀的函數，單位是年。

and it grows to a size of about 10 or 12 thousand ants when the colony is five.

所以蟻群在蟻后建立之初，並沒有任何其它的螞蟻，

And it stays that size until the queen dies

但在五年後，它會成為一萬到一萬兩千隻螞蟻的蟻群，

and there's nobody to make more ants, when she's about 15 or 20 years old.

並且在蟻后死去之前，一直維持這樣的數量規模，

And it's when they reach this stable size, in numbers of ants,

蟻后約在15到20歲時便停止生產。

that they start to reproduce.

而當蟻群的數量達到穩定的大小時，

That is, to send more winged queens and males to that year's mating flight.

它就會開始複製新的蟻群，

And I know how colony size changes as a function of colony age

也就是一年一度送出有翅的新蟻后與公蟻，讓他們飛到外面交配。

because I've dug up colonies of known age and counted all the ants. (Laughter)

而我之所以敢說蟻群的大小會隨著蟻群年紀變化，

So that's not the most fun part of this research, although it's interesting.

是因為我挖開過每一個已知歲數的蟻群，還數過所有螞蟻的數量，

(Laughter)

這還不算是這研究最好玩的部分，不過是蠻很有趣的啦。

Really the question that I think about with these ants is what I call task allocation.

(笑聲)

That's not just how is the colony organized,

其實我真正想從這些螞蟻身上了解的是"任務分配"的問題，

but how does it change what it's doing?

這不僅僅是蟻群如何組織的問題，

How is it that the colony manages to adjust

而是蟻群如何改變自己的行為?

the numbers of workers performing each task as conditions change?

當狀況有變時，他們如何

So, things happen to an ant colony.

調派足夠的工蟻去執行各項工作？

When it rains in the summer, it floods in the desert.

蟻群可能會碰到下面這些狀況：

There's a lot of damage to the nest,

夏天的雨水會淹沒這片沙漠，

and extra ants are needed to clean up that mess.

而重創蟻巢，

When extra food becomes available --

那麼就需要多派些螞蟻來災後重建；

and this is what everybody knows about picnics --

而當有螞蟻發現新的食物，

then extra ants are allocated to collect the food.

每隻螞蟻都會知道野餐時間到了，

So, with nobody telling anybody what to do, how is it that

那麼就要多分派些螞蟻去搬回這些食物。

the colony manages to adjust the numbers of workers performing each task?

所以，在沒有任何人指揮的情況下，

And that's the process that I call task allocation.

蟻群是如何做到派遣足夠的工蟻去執行各項任務的呢?

And in harvester ants, I divide the tasks

而這就是我所謂的"任務分配"問題。

of the ants I see just outside the nest

我將這種收割蟻所做的可能任務，

into these four categories: where an ant is foraging,

依照我在巢穴外觀查到的，

when it's out along the foraging trail, searching for food or bringing food back.

分成四大類，首先是覓食分隊，

The patrollers -- that's supposed to be a magnifying glass --

他們的任務就是沿著覓食路徑出去找食物，並從原路帶食物回來。

are an interesting group that go out early in the morning

再來是巡邏分隊--他們應該需要放大鏡--

before the foragers are active.

他們是很有趣的分隊，每天一大早

They somehow choose the direction that the foragers will go,

在覓食分隊行動前，他們就出門了。

and by coming back -- just by making it back --

不知他們是怎麼選擇方向的，

they tell the foragers that it's safe to go out.

不過他們來回的路徑，就是之後覓食分隊的覓食路徑，

Then the nest maintenance workers work inside the nest,

他們會告訴覓食分隊這些路是安全的。

and I wanted to say that the nests look a lot like Bill Lishman's house.

再來是在蟻巢內工作的修護分隊，

That is, that there are chambers inside,

我想說的是通常蟻巢看起就跟比爾.里斯曼的房子差不多，

they line the walls of the chambers with moist soil

也就是說裡面都是房間，

and it dries to a kind of an adobe-like surface in it.

他們用潮濕的土壤來造隔間的牆，

It also looks very similar to some of the cave dwellings

牆乾了就會形成像泥磚般的表面。

of the Hopi people that are in that area.

蟻巢看起來也很像某些

And the nest maintenance workers do that inside the nest,

當地印地安人的洞穴居。

and then they come out of the nest carrying bits of dry soil in their mandibles.

此外修護工人們在造牆時，

So you see the nest maintenance workers come out with a bit of sand,

他們也會用大顎搬一些乾掉的的泥土到巢穴外，

put it down, turn around, and go back in.

所以你可以看到修護工人夾著一些沙出來，

And finally, the midden workers put some kind of territorial chemical in the garbage.

放在地上，轉個身，然後又回去洞穴。

So what you see the midden workers doing is making a pile of refuse.

最後是堆肥分隊，他們會把當地取得的某種化學物質放在垃圾中，

On one day, it'll all be here, and then the next day

你會看到堆肥分隊不斷地在堆放廢棄物。

they'll move it over there, and then they'll move it back.

可能今天他們把廢棄物堆在這裡，

So that's what the midden workers do.

隔天把它移到另一個地方，然後哪一天又把它們移回來，

And these four groups are just the ants outside the nest.

這就是堆肥分隊的工作。

So that's only about 25 percent of the colony, and they're the oldest ants.

這四個分隊都是需要走出巢穴工作的螞蟻，

So, an ant starts out somewhere near the queen.

他們只佔了整個蟻群的四分之一，但他們是年紀最大的螞蟻。

And when we dig up nests we find they're about as deep

每隻螞蟻都是從蟻后的身邊成長離開，

as the colony is wide, so about a meter deep for the big old nests.

在我們挖了一些蟻巢後，才發現它們的深度

And then there's another long tunnel and a chamber, where we often find the queen,

跟它們的寬度差不多，也就是說一個大的老巢大約有一公尺深，

after eight hours of hacking away at the rock with pickaxes.

裡面有個很長的地道，底部有個房間，我們常常可以在那裡找到蟻后，

I don't think that chamber has evolved because of me and my backhoe

這大約需要用鶴嘴鋤在土石裡挖八個小時。

and my crew of students with pickaxes,

我不認為那個房間會因為我的挖土機或我學生的鶴嘴鋤，

but instead because when there's flooding,

就會有所遷移；

occasionally the colony has to go down deep.

但是當水災發生時，

So there's this whole network of chambers.

蟻穴偶爾會繼續往更深處挖。

The queen's in there somewhere; she just lays eggs.

這是整個蟻穴的地下網絡，

There's the larvae, and they consume most of the food.

蟻后就在那裡某處生蛋；

And this is true of most ants --

那邊是幼蟲，大部分的食物都是用來餵養他們。

that the ants you see walking around don't do much eating.

可以說大部分你平常看到

They bring it back and feed it to the larvae.

在那走來走去的螞蟻，其實是吃很少的，

When the foragers come in with food, they just drop it into the upper chamber,

他們主要是把食物帶回去供養那些幼蟲。

and other ants come up from below, get the food,

當覓食分隊帶著食物回來時，他們通常只是把它放到較上層的存放室，

bring it back, husk the seeds, and pile them up.

然後下層的螞蟻就會上來搬這些食物，

There are nest maintenance workers working throughout the nest.

把它搬回去後，剝去種子的外殼，並堆放好。

And curiously, and interestingly, it looks as though at any time

那邊都是蟻巢的維護分隊，他們遍布整個蟻巢。

about half the ants in the colony are just doing nothing.

令人好奇且有趣的是，蟻群似乎總是有

So, despite what it says in the Bible,

大約一半的螞蟻是不做任何事的，

about, you know, "Look to the ant, thou sluggard,"

所以儘管聖經提到說：

in fact, you could think of those ants as reserves.

"懶鬼啊，看看螞蟻是多麼的勤勞"，

That is to say, if something happened -- and I've never seen anything like this happen,

事實上，你也許會想說他們可能是後備部隊，

but I've only been looking for 20 years --

也就是說，如果發生了某事--不過我倒是從沒看過這"某事"發生過，

if something happened, they might all come out if they were needed.

儘管我只觀察它們二十年 --

But in fact, mostly they're just hanging around in there.

如果發生了某事，或許他們會傾巢而出也不一定。

And I think it's a very interesting question --

但事實上，大部分的時候他們仍是逗留在蟻巢裡，

what is there about the way the colony is organized

我覺得這是一個非常有趣的問題 --

that might give some function to a reserve of ants who are doing nothing?

蟻群為什麼會是那樣的組織方式？

And they sort of stand as a buffer in between

那些做為後備部隊不做任何事的螞蟻，他們扮演了什麼角色?

the ants working deep inside the nest and the ants working outside.

他們有點像介於在巢穴深處的工蟻

And if you mark ants that are working outside, and dig up a colony,

與在巢外的工蟻之間的緩衝角色，

you never see them deep down.

如果你把那些在巢外的工蟻做上記號，然後挖開蟻巢，

So what's happening is that the ants work inside the nest when they're younger.

在蟻巢深處，你是找不到一隻螞蟻是有記號的。

They somehow get into this reserve.

所以其實只有年輕的螞蟻會在蟻巢內工作，

And then eventually they get recruited to join this exterior workforce.

他們以某種方式進入後備部隊，

And once they belong to the ants that work outside, they never go back down.

最後他們會被徵募並加入外面的工蟻；

Now ants -- most ants, including these, don't see very well.

一旦他們被歸屬為外部的工蟻，他們就再也不會回到蟻巢的下層。

They have eyes, they can distinguish between light and dark,

大部分的螞蟻，包括這些，他們的視力不怎麼好，

but they mostly work by smell.

雖然他們有眼睛能夠區分明暗，

So just to reinforce that what you might have thought

但是大部分他們依靠的是嗅覺。

about ant queens isn't true --

所以我想再強調，你們過去對蟻后的印象

you know, even if the queen did have the intelligence

很可能是不正確，

to send chemical messages through this whole network of chambers

即使蟻后是真的有智慧，

to tell the ants outside what to do,

能夠經由巢穴中的內部網路，傳送化學訊息

there is no way that such messages could make it in time to see

去告訴所有外部的工蟻該做什麼，

the shifts in the allocation of workers that we actually see outside the nest.

但這些訊息也無法使那些工蟻像我們實際在巢外看到的那樣，

So that's one way that we know the queen isn't directing the behavior of the colony.

能夠迅速完成工作分派的調動與更替，

So when I first set out to work on task allocation,

所以這是我們知道蟻后無法指揮整個蟻群的原因。

my first question was, "What's the relationship

所以當我開始研究它們的"任務分配"時，

between the ants doing different tasks?

我第一個問題是"不同任務的分隊之間，

Does it matter to the foragers what the nest maintenance workers are doing?

是否存在著某種關係?"

Does it matter to the midden workers what the patrollers are doing?"

修護分隊做的工作對覓食分隊有重要的意義嗎?

And I was working in the context of a view of ant colonies in which each ant

巡邏分隊做的工作對堆肥分隊有重要的影響嗎?

was somehow dedicated to its task from birth

而就我過去所知的觀點，每隻螞蟻

and sort of performed independently of the others,

似乎從出生就一直致力於它的任務，

knowing its place on the assembly line.

他們似乎明確知道他們的工作崗位在哪，

And instead I wanted to ask, "How are the different task groups interdependent?"

並且能夠獨立執行。

So I did experiments where I changed one thing.

不過我想問的是，"不同任務的分隊之間是如何相互依賴的?"

So for example, I created more work for the nest maintenance workers

所以我做了一些小實驗，

by putting out a pile of toothpicks near the nest entrance,

比如說我在一大早，第一批維護分隊尚未行動時，

early in the morning when the nest maintenance workers are first active.

就放了一堆牙籤在蟻穴的入口附近，

This is what it looks like about 20 minutes later.

以便增加維護分隊的工作份量。

Here it is about 40 minutes later.

這是二十分鐘後的狀況，

And the nest maintenance workers just take all the toothpicks

而這是四十分鐘後的狀況，

to the outer edge of the nest mound and leave them there.

最後維護分隊把所有的牙籤

And what I wanted to know was, "OK,

都堆放到巢穴的邊界去了。

here's a situation where extra nest maintenance workers were recruited --

而我想知道的是，

is this going to have any effect on the workers performing other tasks?"

在這種情形下，維護分隊要招募更多的工蟻來支援，

Then we repeated all those experiments with the ants marked.

而這會影響到其它分隊的工蟻嗎?

So here's some blue nest maintenance workers.

然後我們重覆這個實驗，並且將工蟻們作上記號，

And lately we've gotten more sophisticated

我們把維護分隊標記為藍色。

and we have this three-color system.

接著我們再搞得複雜一些，

And we can mark them individually so we know which ant is which.

用三種顏色將工蟻

We started out with model airplane paint

一隻隻的做標記，如此我們可以知道哪隻是哪隻。

and then we found these wonderful little Japanese markers,

我們一開始是用模型飛機的油漆，

and they work really well.

但後來我們發現日本的小麥克筆，

And so just to summarize the result,

非常適合用來做標記。

well it turns out that yes, the different tasks are interdependent.

我簡述一下我們的實驗結果，

So, if I change the numbers performing one task,

呃..沒錯，結果就是不同分隊的工蟻是彼此合作的，

it changes the numbers performing another.

也就是說如果我改變了一個分隊的工作份量，