'Niches' and Adaptive Innovation
It's interesting to think of this article about why evolution progresses at different rates both over time and across regions in terms of how competition among firms for survival drives innovation in a competitive marketplace. The term 'niche' as used below would be, for example, an industry where firms are earning greater than normal profits, or a profitable market opportunity that has not yet been exploited:
No Vacancy, No Evolution, Evolution 101, by Olivia Judson, Commentary, NY Times: Yesterday, I claimed that a major reason large evolutionary changes often don’t happen is that competition from the creatures around you stops you from changing. In other words, in environments that are already rich in different species, natural selection often prevents large changes. My piece of evidence for this was a claim that, when you take other organisms away — when you reduce competition for food, or space — evolution explodes. Today, I want to examine the truth of these claims more closely.
I want to start with a question: Why aren’t there more insects in the sea? Insects, after all, make up the bulk of all known animal species — most animals are insects — yet hardly any insects live in the sea.
It’s not because of the water. Many insects live in freshwater for at least part of their lives. Think of mayfly nymphs, or caddis fly larvae... Or think of water striders, skating along the top of a pond... Could it be the salt? No. Brine fly larvae live in the Utah’s Great Salt Lake. Some mosquito larvae do fine in salt, too — they can even live in the hypersalty Dead Sea. ...
I put it to you that the reason so few insects get beyond rock pools or dip their toes beneath the low tide mark is that the niches they would occupy in the ocean proper are taken already. Crabs, lobsters, shrimp, barnacles, water fleas (which are ... little crustaceans) and company have got all niches covered. My prediction — which I hope we won’t be testing — is that if all the crustaceans were wiped out of the oceans, insects would move straight in.
My prediction rests on the fact that it is typically difficult to evolve to occupy a niche that is already full. An invader must be better at exploiting the niche than the current occupant, who has already evolved to make effective use of it. By contrast, when a niche is empty — when seeds are falling to the ground and no one is eating them, say — it doesn’t matter if, at first, an animal is a bit inept at finding and opening the seeds.
Consistent with this idea is the observation that when new niches open up — perhaps because new islands or lakes or cave systems have formed, or because an asteroid has hit the earth and eradicated millions of species — the first organisms to become established in the new environments evolve quickly and reliably into all sorts of new species. This phenomenon is known as adaptive radiation.
Newly erupted islands are famous for this. Over and over again, archipelagos see explosive bursts of evolutionary change and the rapid appearance of species found nowhere else. New Zealand is full (and was fuller) of an amazing array of unique flightless birds... Hawaii has an abundance of unique fruit flies, spiders, silverswords ... and birds. Madagascar has all manner of lemurs... And everyone knows about the Galápagos.
Rapid bursts of evolution can also happen in new lakes... Indeed, right now, the great lakes of tropical Africa are the backdrop for the fastest known radiation of vertebrates, the cichlid fishes. Lake Victoria, for example, appears to have dried up and then refilled around 14,600 years ago. Since then, perhaps 500 different species of cichlid have evolved... Lake Victoria has cichlids that eat algae, cichlids that eat other cichlids, cichlids that eat fish eggs — cichlids, in short, that have evolved to eat everything that can be eaten. Some fish live in shallow water; others prefer the deeps...
Ideas about adaptive radiation can also be tested in experiments. ...[M]any bacteria can whiz through hundreds of generations in a month. This makes it relatively easy to use bacteria to look at radiations. Here’s what you do. You create two sets of environments, one simple, and one complex. The complex environment might have several different places to live, or a variety of sources of carbon. The simple environment has just one habitat or foodstuff. Then, since bacteria reproduce asexually, you take genetically identical individuals, and release them into the two different environments. Sure enough, mutations happen, and the bacteria rapidly evolve to exploit the different niches. After a month, you will find that bacteria from the complicated environment have become genetically diverse. Those from the simple environment, in contrast, remain unevolved.
In short, empty niches are a license for evolutionary change. Once the new niches are full, natural selection acts to stop further change, and the rate of evolutionary change slows. Fossils, islands and test tubes — they all show the same dynamics. ...
This impliess that it is not competition per se that produces rapid innovation, but competition coupled with emergent profitable 'niches.'
Posted by Mark Thoma on Wednesday, June 21, 2006 at 03:51 PM in Economics, Science, Technology | Permalink | TrackBack (0) | Comments (18)

Ah, thank you :) Olivia Judson series of essays are stunning from beginning to this day. They get me invited to lunch each day.
Posted by: anne | Link to comment | Jun 21, 2006 at 05:36 PM
http://query.nytimes.com/gst/fullpage.html?sec=health&res=9C03EED6153EF936A35752C1A9649C8B63
November 5, 2002
Dr. Tatiana, a Dr. Ruth With Advice for Other Species
By NICHOLAS WADE
DR. TATIANA'S SEX ADVICE TO ALL CREATION
By Dr. Olivia Judson
''Dear Dr. Tatiana,
My name's Twiggy, and I'm a stick insect. It's with great embarrassment that I write to you while copulating, but my mate and I have been copulating for 10 weeks already. I'm bored out of my skull, yet he shows no sign of flagging. . . . How can I get him to quit?
Sick of Sex in India.''
Dr. Tatiana is the pen name of Dr. Judson, an evolutionary biologist who has discovered that an advice columnist's pose is a deft device for discussing the mating game's many amazing variations in nature.
In the replies to her finned, furred and feathered correspondents, titled ''Dr. Tatiana's Sex Advice to All Creation,'' she counsels creatures caught in the most curious predicaments, from a fig wasp in Ribeirão Preto, Brazil, concerned that her lovers keep decapitating one another, to an elephant in Amboseli in Kenya bothered that his penis had turned green, to a male bronze-winged jacana bird of Tamil Nadu, India, cruelly neglected by the female who keeps him in her harem.
Beneath these racy tales and the saucy advice offered for their solutions, a less visible agenda is at work. The reader is nudged to start thinking like an evolutionary biologist. Why did this bizarre behavior evolve and what advantage does it confer? Does the male stick insect copulate for 10 weeks because he has nothing better to do with his time or could it be to prevent other males from fertilizing his inamorata's eggs?
Eggs are few, and sperm are many. This microscopic-level asymmetry is the root cause of ardent civil war that in every species pits male against male, and male against female. Males, from sea lions and fruit bats to to Taliban, are driven to control females' fertility so as to ensure their own paternity.
But a female's interest usually lies in having many lovers. There are many sound reasons for the female to pursue a policy of determined promiscuity, Dr. Tatiana informs a male yellow dung fly of 12, including to guard against male sterility, to ensure diversity in her offspring, to encourage each male in her group to think that he is the father and protect her children accordingly and to encourage competition among the sperm of several males so as to ensure her egg gets the best.
''Natural selection, it seems, often smiles on strumpets,'' Dr. Tatiana writes without much hint of regret. ''Sorry, boys.''
Despite Dr. Tatiana's dark and worldly wisdom, her creator is a fair-haired young woman with an infectious laugh. Dr. Judson studied biology at Stanford and at the University of Oxford in England, where she was one of the last students of Dr. William D. Hamilton, whose theory of kin selection undergirds large arenas of modern biology, including sociobiology and evolutionary psychology. In 2000, he died of malaria that he caught after a trip to Congo to investigate a hypothesis about the origin of AIDS.
In an interview, Dr. Judson recalled her mentor's imaginative gift for entering the minds of the insects he studied.
Once she asked him about the biology of a species of asexual weevil. A few days later, a pinned specimen arrived in her mailbox.
But there was the occasional disagreement. ''I once had an argument with him,'' she said. ''I burst into floods of tears, and he sat staring at his toes.''
Dr. Hamilton was famously impractical. University administrators required him to give just one lecture a year. But on at least one occasion he forgot even that. As a profound evolutionary thinker, Dr. Hamilton knew the difference between a sex symbol and the real thing....
Posted by: anne | Link to comment | Jun 21, 2006 at 05:38 PM
Hmmm, what was it that Dr. Hamilton knew about a symbol and the real thing :)
Posted by: anne | Link to comment | Jun 21, 2006 at 05:44 PM
I always thought that Schumpeter's 'creative destruction' was at least partially misunderstood & misreported by the mainstream business media. They report it sort of as the forces wrecking the economy create the new growth.
I see it as a two phase process... subtle difference but important. The destructive forces clear out the existing competition (old technology) and open up niches that remain less competitive for atleast awhile (until fully populated). So you would need TWO simultaneous factors (1) niche clearing and (2) young innovative firms to populate those niches.
The policy implications ARE very different... I don't see gov't eagerly expediting 'niche clearing' even as they try to nurture new young firms... clearing niches is pretty painful. Infact I see them doing the opposite. But nurturing small enterprise is pretty useless WITHOUT space for them to grow. These two objectives - protect the old & nurture the new - are counter each other.
Also I'm not keen on the 'evolution' analogy. I think a better analogy is forest cycling... Forests mature and as they do they crowd & shade out the underbrush & saplings beneath them... even their own offspring.
In fact many species won't even germinate until the cone or seed coat has been burned by fire. And as we know fires are frequent events in forests though few actually crown & consume the mature trees. In these routine fires saplings and brush get consumed... new ones sprout & are likewise consumed again later in the future.
As these forest age they get sicker & eventually enough fallen trees, duff & material builds up on the forest that when a fire comes through (often coupled with an environmental change like a drought) the whole forest burns... fire 'crowns' & kills even the largest mature trees.
What is left looks like a disaster - except for the sees that will be the next generation of saplings - to them it now looks like *HEAVEN ON EARTH*. They sprout. They grow. They battle each other for sunlight, water & nutrients... start on their path to becoming mature trees... and most never make it.
But without the 'forest clearing event' which required an old and aging forest and/or a severe drought which allowed what would have been just a routine fire to turn into a 'fire of the century'... none of those new saplings would have had a chance to mature.
Analogous factors...
Mature forest, trees = mature industry, companies
Saplings = innovative smaller firms
Routine Fires = Normal business cycle recessions
Drought = Economic pressure from shifts in technology or economic environment
Duff = Corruption, subsidies, protectionism
Fire of the Century = Depression
Analogies are never perfect but this one helps me make sense of it all,even if imperfect... but then I started out in biology even before I went into engineering which was LONG before I ever started thinking about economics... so it works for me. But then I'm a moron - ask my kids.
:::::
BTW - I think our 'forest' is getting pretty mature again... I believe we are 'past peak' in the IT/Telecomm wave started in the mid to late 80s... peaked with the internet economy around 2000 and our mature trees & forests are now aging pretty rapidly (decelerating innovation, commoditization of product offerings, consolidation of industries) but are still a ways off from a 'crowning firestorm'. However a 'severe drought' could accelerate that process.
Posted by: dryfly | Link to comment | Jun 21, 2006 at 09:30 PM
«This impliess that it is not competition per se that produces rapid innovation, but competition coupled with emergent profitable 'niches.'»
In the tech world this is sort of common knowledge: displacing well entrenched incumbents is impossible or very expensive, and the cost must be born by cross subsidy (e.g. Microsoft using the OS profits to grab Wordperfect's office suite market).
Hey, IBM is still the incumbent mainframe monopoly like in the 1960s, nobody managed to displace them. More like Microsoft sidestepped them.
Posted by: Blissex | Link to comment | Jun 21, 2006 at 10:12 PM
http://judson.blogs.nytimes.com/?p=49
June 19, 2006
Waiting to Evolve
By Olivia Judson
Yesterday, I mentioned that one of the big questions of modern evolutionary biology is: Why hasn't there been more evolution? Today, I want to look at this in detail.
I'll start by putting it into perspective. For Darwin, a major worry was being able to persuade people that enough time had passed for the evolutionary changes that he documented to have happened. But now that we can look at genes and measure natural selection directly, it is obvious, as I showed yesterday, that it can happen very fast. Yet, it often doesn't seem to happen at all. In particular, as Stephen Jay Gould, the late paleontologist and writer, pointed out, when you look at the fossil record, you have the impression that forms tend to stay more or less the same for long periods of time. In other words, there seems to be a mismatch between the apparent stability of the fossil record and the evolution that we actually observe today. Why is this?
One reason we often don't notice big changes has to do with the fact that natural selection acts in different ways on different genes. Now that we can analyze genes, we can see where natural selection actively stops changes from happening, and where it pushes for rapid change. Interestingly, many of the most rapid changes happen in genes that affect traits we can't see with our eyes. For instance, we've discovered that, in mammals, two types of genes are especially prone to changing fast: genes of the immune system and genes involved in making sperm. But the products of these genes — the chemicals a sperm wears on its head and the weapons the body uses to fight diseases — leave no fossil trace.
But that is not the only reason we sometimes don't notice evolutionary changes. A variety of forces can keep a population doing the evolutionary equivalent of treading water. The simplest is if natural selection is fickle, gusting this way and that, continually changing direction. Look no further than the famous finches of the Galápagos Islands (evolutionary celebrities, because they helped inspire Darwin to develop the idea that evolution happens because of natural selection). A 30-year study of the medium ground finch, Geospiza fortis, has shown that the best beak size shifts from year to year — and if you measure beaks carefully, you can watch the average size shift about. In droughts, a big beak is an advantage, because you can crack open tough seeds more easily. In wet years, when small seeds are abundant, having a smaller beak is better.
If this sort of fickleness is typical — and it may well be — we wouldn't expect to often see much significant change. Indeed, we can make a prediction: to see big changes, natural selection must blow in the same direction for a long time....
Posted by: anne | Link to comment | Jun 22, 2006 at 02:55 AM
Hey, everybody should read some Jane Jacobs - encouraging ecomic diversity is the way to go. Why are Stanford and MIT so successful at stimulating innovation? How should we stop IP from stifling innovation and co-operative research. More research needed.
Posted by: reason | Link to comment | Jun 22, 2006 at 03:06 AM
This form of competition is behind several theories that try to explain the long-wave in economic history.
You have a new technology that leads to a wave of
innovation and rapid growth. As that waves matures
the growth slows and there is no new source of
technology large enough to provide strong growth. So overall growth slows and will not reaccelerate until some new technology generates a new large scale wave of technology.
Within this wave you see a pattern of many relative small startup type firms in the early stages. But
over time these firms consolidate so by the late stages of the wave you are left with a few large firms that have evolve a relatively stable form of competition.
I think there is a lot to this thesis even if it has been neglected by main stream economics.
Posted by: spencer | Link to comment | Jun 22, 2006 at 05:57 AM
Every city with a sick economy is looking for the illusive high tech niche.
In the meantime they are finding other niches - such as "logistics management" (that is a fancy term for running warehouses for moving Chinese-made merchandise - jobs are low paying).
Posted by: save_the_rustbelt | Link to comment | Jun 22, 2006 at 09:51 AM
..and what dryfly said - good analysis
Posted by: save_the_rustbelt | Link to comment | Jun 22, 2006 at 09:58 AM
Olivia Judson is an excellent writer with a fascinating subject. Thanks for posting this article!
Posted by: Holly W. | Link to comment | Jun 22, 2006 at 11:12 AM
I love this woman :)
http://judson.blogs.nytimes.com/?p=51
June 21, 2006
Use It or Lose It
By Olivia Judson
One of the things I enjoy about studying evolution is that it allows us to make powerful predictions about the natural world. This is because evolution often follows predictable paths. Today I want to talk about one of the most predictable phenomena of all: the evolution of loss or, as it was once called, degeneration.
I’ll start, as I often do, with a question. Why do so many of the animals that live in the deep sea have eyes? At first, this seems extremely odd. The sun doesn’t shine in the vasty deep — it’s an ocean of blackness. Animals that live in dark caves, be they millipedes, beetles or fish, reliably lose their eyes and their colors. Why doesn’t this happen in the sea?
Why do I ask? Well, here is a general law of nature: When organisms no longer use something they have — whether we’re talking eyes, brilliant colors, wings or even genes for making energy from sunlight — that something rapidly evolves to become lost. As I mentioned yesterday, birds living on remote islands routinely lose the power of flight. Fruit flies that evolve to be asexual — reproducing by laying eggs that don’t need to be fertilized — fast lose the ability to respond to male advances. The soil bacterium Myxococcus xanthus normally has (for a bacterium) an elaborate social life; bacteria hunt together and cooperate to build spores. But if you keep the bacteria in bottles of liquid, you can put an end to their social lives by continually shaking the bottles and interrupting their social discourse. After generations of such treatment, the bacteria evolve antisociality: they just can’t work together anymore.
And — this is especially striking — full-time parasites lose most of their bodies. They also lose many of their genes. Microsporidians, which are strange single-celled fungi that are parasites of animals, have lost their mitochondria. (Mitochondria are what cells use to make energy from sugar.) Animals that become parasites can even lose their brains. This is how one early evolutionary biologist, Edwin Ray Lankester, put it in 1880:
"Any new set of conditions occurring to an animal which render its food and safety very easily attained, seem to lead as a rule to Degeneration. … Let the parasitic life once be secured, and away go legs, jaws, eyes, and ears; the active, highly-gifted crab, insect, or annelid may become a mere sac, absorbing nourishment and laying eggs."
There are two roads to loss. The first is via something known as “relaxed selection.” The idea is simple: If a given trait — say, fin color — no longer affects how well you can survive or reproduce, mutations accumulate in the genes for fin color, thus destroying the ability of those genes to work.
Relaxed selection is easy to show in a laboratory. Suppose you have a vial full of fruit flies. You assign a mate to each fly at random — thus rendering irrelevant the ability to seduce — and then choose two offspring from each pair to put into the next generation. This way everyone has the same number of surviving children (which means, as you may recall from one of my earlier columns, that natural selection cannot operate). Furthermore, you give everybody a life of luxury and ease, so finding food isn’t a problem. After repeating this for some number of generations, you make life difficult again, and see what happens. One such experiment showed that after just 30 generations, a fly’s ability to survive in difficult conditions had dropped sharply.
If selection is relaxed in this fashion for a long time, the rate of degeneration may start to accelerate. This is because stopping mutations from happening is itself something that usually falls under natural selection. If there is no penalty for having more mutations, more mutations will start to happen.
The second road to loss is when natural selection actively drives it. This is called selected loss. Then, it’s not simply that eyes (or whatever) no longer help you; they actually get in your way. For instance, for many parasites, there is a premium on being able to reproduce quickly. This means that losing genes you don’t need anymore is actually an advantage because you can copy your genome faster if it is smaller. Or, consider flight in birds....
Posted by: anne | Link to comment | Jun 22, 2006 at 12:31 PM
«Why are Stanford and MIT so successful at stimulating innovation?»
As compared say to IIT or Tsinghua? :-)
Perhaps (heavy handwaving here) because there are VCs queueing to throw money at any Stanford or MIT (or Oxford or Cambridge) start-up because they come from such prestigious places and because they are next door and because the USA has some much free money and capital sloshing around that it can fund zillions of stupid start-ups until a Sun or Google pop up?
The number of failed innovations from Stanford or MIT (or Oxford or Cambridge) is quite awesome, but people see only the successes, and miss out the ratio between resources invested and number of successes.
Now the question is whether throwing the same wall of money at Oregon or Miami or IIT or Tsinghua start-ups would result in a better resources/successes ratio, but the answer is not going to happen anytime soon.
Posted by: Blissex | Link to comment | Jun 22, 2006 at 12:33 PM
"Any new set of conditions occurring to an animal which render its food and safety very easily attained, seem to lead as a rule to Degeneration. … Let the parasitic life once be secured, and away go legs, jaws, eyes, and ears; the active, highly-gifted crab, insect, or annelid may become a mere sac, absorbing nourishment and laying eggs."
Humans are some sort of animals, right? So, can this happen to me? I'm hoping that with high gas prices I'm going to get a bike and my legs are not going to go so soon...
Posted by: | Link to comment | Jun 22, 2006 at 12:59 PM
http://judson.blogs.nytimes.com/?p=35
May 31, 2006
Genie, I Dream of Planets
By Olivia Judson
Call me a geek, but if a genie appeared before me and offered me a wish, here's what I'd wish for: detailed descriptions of at least 127 other randomly chosen planets in the galaxy, of which 63 would be home to life.
This may sound greedy (wishes usually do). After all, life on earth holds uncountable mysteries. Here's one that puzzles me: Why is it that no birds have evolved pregnancy? Representatives of all the other major groups of backboned animals, from guppies to skinks, have evolved to give birth — why not birds? It would surely be better for an emperor penguin to give birth than to play ice hockey with its egg. Another: Can you alter the odds of conceiving a son or a daughter by altering how often and when you have sex? Tentative evidence suggests you might be able to — but the data are deeply unclear. And how about this: What, apart from thinking, does the sperm whale use its head for? A sperm whale's head makes up about one third of its body length, and much of the space is taken up by something called the spermaceti organ — but despite much speculation (is it acoustical? A battering ram?), no one knows exactly what this does.
Yet for all the intriguing questions on earth, with only one planet to study, it's impossible to know which features of life are general features, and which are unique to the earth. We can't even be sure what it is about this planet that makes it good at sustaining life: is it the oceans? The motions of the continents? Its magnetic field? Its size? Its moon? Its distance from the sun? Our general situation in a distant and unfashionable section of the galaxy? All of the above? (All have been invoked.) None?
When a volcano erupts in the ocean and forms a new island, at first, there is no life. But before long, organisms arrive. What happens when a new planet forms? Does life evolve de novo each time? Or does it arrive, by comet or by meteorite? Once life gets going, what happens then? On earth, organisms alter their local environment — and in doing so, open up niches that allow other organisms to flourish. For instance, think of reed beds around a marsh. Over time, the reeds make land: mud and silt collect around their stems, and when they die, their corpses help clog up the water's edges. Eventually, the lake may disappear; then, the reeds will have sculpted the landscape so drastically they can no longer live there.
Do organisms wreak such changes on the scale of an entire planet? Some evidence suggests they do. We know, for instance, that oxygen was not a significant feature of the earth's early atmosphere; instead, it has built up over the eons, a deadly waste product given off by organisms that gather energy from the sun. As oxygen levels have risen, the sorts of creatures living here have changed. It's even been suggested that the metabolic activities of early organisms helped the formation of the first continents, by altering the chemical composition of rocks. Perhaps drifting continents are an inevitable consequence of life. If we had more planets, we could know.
If we had more planets we could also probe the pace of evolution. On Earth, evolution proceeds surprisingly slowly — yes, slowly....
Posted by: anne | Link to comment | Jun 22, 2006 at 01:23 PM
The beginning of Olivia Judson's essays was on May 28, and one is better than the other and all are wonderful and have all sorts of analogies laden in them as biology always will for biology is of "us."
Posted by: anne | Link to comment | Jun 22, 2006 at 01:25 PM
Mark Thoma has done us a singular favor by attending to biology and evolution in posts. I have long complained, and am being taken seriously about the use of analogies from physics and chemistry rather than biology and psychology in social studies analysis; notice, I term economics as a social study as does our college though our President may object :)
Posted by: anne | Link to comment | Jun 22, 2006 at 03:43 PM
Life and thoughts of life are where our analogies should come from in the social studies as in philosophy. There is all sorts of comment going about on Olivia Judson's essays :)
Posted by: anne | Link to comment | Jun 22, 2006 at 03:45 PM