| 1 | DARWINâS ARGUMENT AND THREE PROBLEMS: HERITABILITY, SEXUAL SELECTION AND ALTRUISM |
Some of the questions addressed in this chapter:
- What is natural selection?
- What problems did the theory of natural selection face after Darwinâs death?
- How were the problems addressed?
- What are the laws of inheritance?
- What is sexual selection?
- How can we account for altruistic behaviour according to natural selection?
SOME KEY TERMS AND CONCEPTS
Adaptation; Altruism; Chromosomes; Darwinism; Fitness; Function; Genes; Heritability; Lamarck(ism); Mendel(ian); Natural selection; Reproductive success; Sexual selection; Variation.
LEARNING OBJECTIVES
Having studied this chapter you should be better able to:
| Outline the theory of natural selection. |
| Indicate which parts of the process Darwin did not fully understand. |
| Outline the mechanisms which underlie evolution. |
INTRODUCTION
In this chapter we will look at Darwinâs argument for the theory of natural selection and its logic. We will see that it is a good theory in that it offers a parsimonious account of the fact that animal and plant species are typically well designed to survive and reproduce in the environments in which they occur. Soon after its presentation, Darwinâs theory gained currency in the scientific community and was widely accepted as a plausible account of how species come about and evolve over time. However, it still faced a number of problems. It wasnât entirely clear how characteristics of parents were passed to offspring, why some characteristics seemingly detrimental to long-term survival persisted in various species, and why all organisms were not relentlessly selfish. These problems were tackled in time and we will briefly review the solutions.
DARWINâS ARGUMENT
BOX 1.1 ARE YOU A DARWINIAN?
Before we begin, ask yourself:
Do you believe that humans have evolved?
Do you accept that that you are the product of evolution?
Do you believe that humans are adapted?
Do you accept that you are fitted to, or designed to survive in, certain specific environments?
Do you believe that what we are adapted to and for can be addressed by our natural history?
Do you believe that the physical form that you take is explicable in terms of past selection pressures?
I suggest that if you either do or are inclined to say âyesâ to these questions then you are indeed a Darwinian.
Charles Darwinâs argument for the natural selection of evolved life forms is not complicated and is based on observations. As was suggested soon after the publication of his On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (Darwin, 1859), Darwinâs interpretation of his evidence appears, on hindsight, to be obvious. The theory of natural selection can be presented as comprising three basic premises: the variation premise, the heritability premise and the adaptation premise. Together these premises led Darwin to the conclusion that life forms on earth evolved by a blind process he called natural selection. Let us look at the premises a little more closely.
VARIATION
The variation premise is based upon the observation that organisms within a species differ in their physical and behavioural characteristics. Following the conventions of common language, Darwin called identifiable differences between individuals of the same species âvariationsâ. Variation is demonstrated by the fact that no two instances of a species are physically (save the possibility of monozygotic siblings, more commonly called âidentical twinsâ) or behaviourally identical.
INHERITANCE
The inheritance premise is based on the observation that variations between members of a species are frequently transmitted from parent to progeny. This is simply to say that identifiable characteristics of individual organisms are passed on to their offspring such that the offspring are distinct from others by virtue of the characteristics. This observation allows us to say that, ceteris paribus (all things being equal), offspring will resemble their progenitors more than they will any other randomly selected member of the species population.
ADAPTATION
The adaptation premise is based on the observation that organisms are âfittedâ to their environments. To say an organism is fitted to its environment is to say that in the wild (as opposed to in a zoo or a laboratory) it exhibits physical and behavioural characteristics which enable it to cope with and exploit features of the environment in which it lives. These features Darwin called âadaptationsâ.
DARWINâS CONCLUSION
Having established that organisms vary, that the variations can be inherited, and that species were adapted to the environments in which they are naturally found, Darwin inferred that not all variations are âequalâ. What he meant by saying they are not equal is that not all individual differences fitted the carrier equally well to the environment and the problem of reproduction. Darwin argued that as a result of variations some members of a species not only survived longer than others but, crucially, some produced more offspring than others. Invoking the observable fact that variations are heritable, Darwin concluded that variations which facilitate survival and reproduction will be more numerous in the next and future generations than others. The continual and inevitable reiteration of the process wherein some members of a species reproduce and pass on the characteristics which enabled them to survive and reproduce is what we call natural selection. The iteration of this process shapes and reshapes a species over time. Such shaping and reshaping is what we call evolution.
DARWINâS PROBLEMS
As straightforward as it appeared to those of his contemporaries such as Thomas Huxley who proselytised on Darwinâs behalf, Darwin himself and critics within the scientific community identified problems with his theory of evolution by natural selection. One of the problems concerned the observable fact that many typical features of different species appeared to hinder rather than help them to survive. The male peacockâs tail is, perhaps, the most salient and oft-given example. Its size and visibility makes it expensive to produce in metabolic terms and renders the bird vulnerable to predators. We will call this the âproblem of non-fitnessâ. Darwin solved the problem himself in his volume The Descent of Man and Selection in Relation to Sex (Darwin, 1871) with his theory of sexual selection but his solution was not widely accepted until the 1930s. A second problem was the mechanism of inheritance. Darwin accepted that he did not know how it was that adaptive variations were transmitted from parent to offspring. He also did not know that the mechanism and rules of inheritance had, to a considerable extent, been discovered by Gregor Mendel. And a third problem, and one that Darwin said made him feel âsickâ, was that of altruism. According to the theory, organisms should behave purely in their own self-interest but innumerable observations suggested that this âruleâ was routinely broken by any number of different species. The solution to the problem of altruism was presented in the 1960s by William Hamilton. The problem of altruism was solved by seeing the correct level at which natural selection operates â the genetic level. Let us now look at the solutions to the problems of non-fitness, heritability and altruism in turn. What we learn here will be essential to our understanding of later analyses of the evolutionary basis of human thought and behaviour.
THE PROBLEM OF NON-FITNESS AND SEXUAL SELECTION THEORY
As we have noted, many organisms, including humans, exhibit physical and behavioural characteristics that are typical of the species but appear to be detrimental to the bearerâs prospects of survival and longevity. An example that we will examine in more detail in Chapter 8 âCompetition, aggression and violenceâ is what has been called the âyoung male syndromeâ â the apparently unnecessarily risky, and often life-threatening, behaviour exhibited by post-pubescent human males (Wilson and Daly, 1997). Being typical of a species any such characteristics are, according to theory, evolved and therefore, naturally selected. But, if the characteristics mitigate against survival and longevity how could they be repeatedly selected for and why would they persist?
In The Descent of Man and Selection in Relation to Sex Darwin reinforced his argument that evolution does not favour longevity per se, but it favours reproductive success. Now, while any given organism has to survive for some period of time â a minimum enough time to reach reproductive maturity â its reproductive success determines how many of its characteristics will be represented in the next generation rather than its life span. Of course, there is a relationship between the two given that longevity is likely to aid reproductive success. However, the maxim âDonât count the candles on the cake, count the kiddiesâ holds. What we now call Darwinian fitness â the long-term survival over evolutionary time of any given heritable characteristics â is determined by the reproductive success of an organism and not the length of its life. In the currency of natural selection it is of no use if an organism lives very much longer than other members of her or his species if he or she does not reproduce.
With this insight in mind let us now outline sexual selection theory. Sexual selection theory argues that physical and behavioural traits that mitigate against longevity â that is, carry a cost to the bearer with regard to overall life expectancy â but facilitate reproduction â that is, aid the bearer in attracting mates and parenting viable offspring â can persist in a population over time. Let us return to the peacockâs tail for an example. While the large and elaborate plumage of the male bird is costly to grow and makes the animal vulnerable to potential predators it also makes it visible and attractive to pea hens. In the case of peacocks the trade-off between the cost to longevity and the gain for reproductive success have favoured the elaborate tail which characterises the birds we see today. In short, sexual selection theory argues that the existence of variable heritable traits which seem to be useless or disadvantageous to survival can be explained if it can be shown that they confer an advantage with respect to reproductive success.
Sexual selection and natural selection are not necessarily mutually exclusive. A trait that enables an organism to enjoy reproductive success may also enable it to live longer. For example, better than average eye sight may enable an organism to spot predators, prey and this ability might make that organism more appealing to members of the opposite sex. However, to be confident that a given trait has evolved by sexual selection an analysis of its function should show that the trait is neutral or detrimental with regard to longevity and that it clearly facilitates reproductive success.
This is the reasoning behind the solution to the apparent problem of non-fitness. When we see that success over evolutionary time is determined by reproduction rather than survival we can also see that physical and behavioural characteristics which appear non-fit in survival terms may be fitness enhancing in reproductive terms.
As has been suggested, Darwinâs proposal was not readily accepted when first introduced and it was further undermined by its rejection by the co-founder of ...