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By Steve Davis | April 9th 2009 11:58 PM | 6 comments | Print | E-mail | Track Comments
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About Steve Davis

gadfly: noun (1) fly that stings horses and cattle. (2) (derog) annoying person, esp one that provokes others into action by criticism, etc.... Full Bio

In works on evolution written by a certain class of biologist we can often see “for the good of the species” references derided in no uncertain terms. Comments such as “fuzzy thinking”, “they got it wrong” and so on have become so habitual that they almost go unnoticed. But is the “for the good of the species” idea really all that bad? It might well be that some comments and discussions are indeed fuzzy, in that they might be poorly thought out or presented. But here’s a discussion from Robert Ardrey’s The Social Contract to consider. Ardrey described the communication between starlings as an element in their defence against peregrine falcons, the falcons being hindered in their attacks by the speed for which they are famous. It turns out that they are so fast that they can only attack prey in the air, as contact with the ground, however slight, would cause injury. This also means that they cannot attack groups of birds; they can only target isolated individuals.

“At such speed, should anything go awry and the falcon strike its prey with a wing instead of its talons, the wing will be broken. And so, through the ages, starling and falcon have perfected their relation to the nicest of balances. Sighting a hovering falcon, the most alert member of the starling flock gives the instant alarm call. Now all crowd together nearly wingtip to wingtip, and begin their unpredictable mass manoeuvres. There is no leader. Somehow all participate in a common sense as to what the next swerve or dip will be. The falcon of course dares not dive. So perfect is the starling social defence that any attack would be suicidal. And so he waits, hovering. And natural selection takes its toll. If one of the hundreds of massed birds has weakness of wing or fails in some fashion to sense the next turn, then it is separated from the flock and the falcon has it. And the unequal starling will leave no further descendents to encumber the flock’s gene pool. Starling defence illuminates the value of society to a group of unequals. In any given situation of danger one member, whether superior in alertness or simply lucky, will be the first to sense threat. And with the alarm call, the perception of the first becomes the property of the group.”

Now Ardrey is clearly talking here of behaviours that he believes are for the good of the group or the species. So is the argument “fuzzy”? Is the pattern of behaviour he describes for the good of the group, or for the good of those individuals who use the group to avoid predation? It may be impossible to say with certainty. Arguments can be put to support both cases. But it is possible to imagine a scenario in which there is a benefit for the flock and ultimately the species. (Selfish gene theorists will be quite at home with this style of thinking – imagined scenarios are their stock-in-trade.)

Imagine a flock of starlings that are preyed upon relentlessly by falcons. After several breeding seasons the starling gene pool may be altered to such an extent that the flock is able to outlast the hovering falcons. At this point the falcons have to make a decision – seek alternative sources of food or starve. Hardly a choice at all. Now unimpeded by constant harassment, the invigorated flock, stronger, faster, smarter, increases in numbers and thereby comes to dominate the starling gene pool. (Keep in mind that the “arms race” argument whereby as the prey becomes more efficient at evasion the predator becomes more efficient at predation, does not always apply. If an alternative food source can be accessed with less effort expended than in hunting starlings, the alternative will become the food of choice.)

Clearly such an outcome, overcoming a predator, is good for the group. Clearly also, while we cannot predict where, when, or how such an outcome will eventuate, processes similar to that which we have imagined have occurred in evolutionary history.

So for critics of “for the good of the species” thinking, it seems it’s all bad news. The concept appears valid. But then, they could have worked that out themselves.

Let’s take our hypothetical flock a little further. Able to outflank predators at every turn, their numbers double, then treble. All is well until famine hits the region. Their food supply drops below the level needed to sustain their greater number. The question now arises, is sudden death by predation as in former times, preferable to a slow death by starvation? And what behaviours will the flock display as it struggles with this new reality? Will they remain a unified whole and migrate to new domains? Will they split into smaller groups and disperse randomly? Will they be forced by the defences of other groups to remain where they are until their numbers drop to sustainable levels? The possible outcomes are countless, but a question keeps nagging in the background. Was their defeat of the falcons truly a victory; was it good for the species in any lasting sense? For that matter, is it possible in evolutionary analysis to even use the words “for the good of”?

There is one thing of which we can be sure. No starling, no matter how fast or strong, can survive alone against its normal range of predators. Of all the options available to deal with a changing environment, the successful option will involve group activity of one type or another. Ernst Mayr once said that there are no laws in animal behaviour, there are only regularities. (Let’s call that Mayr’s Law!) This is one such regularity.




Comments

You have badly misunderstood the issue of levels of selection. Please consult Williams, Dawkins, and others who have written about this at length. Selection occurs at gene, individual, group, species, and clade levels, but not all of these forms of selection are capable of shaping *adaptations*, i.e., devices like falcon claws or behavioral programs for capturing prey. Selection at clade and species levels is entirely incapable of producing directed cumulative changes in gene frequencies (aka evolution) because it wipes out gene pools, rather than modifying them. Group selection IS capable of shaping adaptations, as evolutionary biologists have long recognized (see Williams, 1966), but ONLY under very strict conditions which are rarely met in nature. In most cases group selection is weak compared with selection at gene or individual levels. This does not imply in any way that adaptations will not benefit the group (as suggested in your post), but only that when the good of the individual is at odds with the good of the group, adaptations will be structured to benefit the individual rather than the group. Your example of prey animals flocking is not illuminating because flocking behavior benefits both the individual and the group so this behavior cannot help distinguish which force dominates - individual level or group level selection.

Sex ratio provides a better example. In general, individuals can benefit the group by producing more female offspring than male offspring. This is because females usually invest more in offspring, from producing a larger gamete (egg vs. sperm) to extended care (e.g., internal gestation in mammals). Hence, the number of females is a bottleneck for reproduction and total reproduction would be increased by a population with more females than males. The widespread 50/50 sex ratio shows individual selection overcoming the good of the group/species. Individuals benefit by producing roughly equal numbers of males/females (with some caveats, see Trivers-Willard effect). There are many many other examples of group and individual level selection at odds, and in virtually every case individual selection wins out.

Steve Davis's picture
this behavior cannot help distinguish which force dominates - individual level or group level selection.
Err...that was the whole point of the post. There seems to be a need in some people to see domination when it would be more helpful to see interactions and patterns in biological systems. Thanks for the anonymity too.

Steve Davis's picture
And who's this Dawkins bloke you refer to? Is this the same Dawkins whose understanding of evolution is so flawed that he defines the survival of replicators, an outcome of evolution, as its cause?

Gerhard Adam's picture

"...adaptations will be structured to benefit the individual rather than the group..."

Sorry but this is a chicken/egg argument.  It is obvious that the gene will affect the evolution of the individual, however depending on the state of the group, this will also affect the direction such selection can take.


Reproduction is also a function of what females find attractive.  Therefore natural selection cannot act on the individual without examining the context of that selection on the group.  As the group becomes more important, then the direction natural selection takes will tend to favor individuals that blend with the group.  At this point, it would be extremely difficult to suggest that it is individual selection that is directing the species and virtually everything becomes for "the good of the group".


This shouldn't be taken to mean that the genes are acting on behalf of the group directly, but rather that the genes that will be expressed are only those that attract females.  This is precisely why animals can become social since the group conveys a greater likelihood of survival to the offspring.  Therefore any genetic expression must take into account how well that animal can interact with the prevailing group.  Failure to do so would tend to prevent that gene from entering the gene pool of the group and consequently it would not contribute to the evolutionary direction of the group.


"The widespread 50/50 sex ratio shows individual selection overcoming the good of the group/species."

I'm not sure what this means, because it clearly illustrates how sex ratios promote the group with individuals (potentially) having different strategies to maximize reproductive success.  In other words, there are numerous exceptions to the 50/50 sex ratio within individual families and this ratio doesn't manifest except in groups (since there must be a sufficiently high group interaction to allow for the necessary genetic diversity).  However, using humans as the example, this is misleading since much is dependent on the law of large numbers, since such ratios could not be counted on in smaller tribal societies.  So at that point, we had group interactions (sometimes peaceful, sometimes violent) that broadened the genetic pool.



That's not right. It's not even wrong.

logicman's picture
You have badly misunderstood the issue of levels of selection. Please
consult Williams, Dawkins, and others who have written about this at
length.

I have, Anonymous, and they have badly misunderstood the issue of levels of selection.

There is only one level of selection: the inability of an individual to produce offspring.

Selection is a sieve.  What falls through the holes becomes sterile, effectively sterile, or dies.  The 'sieved out'  DNA is removed from the gene pool.  The population expands again to fill its niche to capacity.  The same, or a new 'sieve' operates, and the cycle repeats.  Over a long enough period of time, a population which once spanned a stream and freely interbred becomes two different species, one each side of an un-navigable torrent.

Of all the options available to deal with a changing environment, the
successful option will involve group activity of one type or another.

And by that activity, enacted not by exact clones but by minutely varying biological individuals, the species remains the sum of its parts at all times, even whilst becoming a new species.

Steve: next time I miss one of your articles, fire up a chat box!  :)

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