Behavioural Interactions

Sexual behaviour

1. Male & Female investment

Male and female investment

Monograph page 41-42

According to the Guinness book of records the largest number of offspring fathered by a single human male is 888, but the record for a human female is 69. Even allowing for some inaccuracy in the numbers there must be a biological explanation for the big difference between the records for mothers and fathers.

Gender in any animal vertebrate or invertebrate is defined simply by the production of sperm by males and eggs (ova) by females. Sexual reproduction occurs with the fusion of a sperm cell with an egg cell.

Parental investment
Monograph page 41-42

There are costs associated with reproduction and these are known as parental investment. The costs of parental investment are not often shared equally between the sexes and they are usually greater in females. The inequality arises during gamete production and investment beyond conception.

 Males produce large numbers of small motile sperm, each with a low individual cost (inexpensive gametes).

head containing
the nucleus which stores the genetic information

Motile tail

length of sperm = 60m

 Males produce large numbers of small motile sperm, each with a low individual cost (inexpensive gametes). Females produced fewer, but much larger eggs, each of which has a nutrient-rich food store and therefore a greater energy cost (costly gametes).
yolky cytoplasm (food store)

nucleus

(sperm on same scale)

Length of ovum = 500m

 Considering the cost of gamete production alone it is clear, in the fertilisation of an egg by a sperm, that the female has made a much bigger parental investment than the male.
This difference has important consequences for the relative costs and benefits of reproductive decisions. It helps to explain conspicuous differences in the behaviour of males and females, such as which sex is more likely to show complex courtship behaviour or parental care.

Parental investment
Parental investment does not stop at gamete production, the concept includes all contributions by a parent that increases the probability that an offspring will survive to reproduce (which may include possible parental care). The costs may also include energy and resources required for courtship and mating, embryonic protection and incubation, and the nurturing of young.
BBC - Sumatran orangutan

External and Internal Fertilisation

Sexual reproduction requires the coordination of behaviour between organisms of different genders. This can mean the simultaneous release of gametes into an aquatic environment in species with external fertilisation, or the sometimes very lengthy courtship which precedes internal fertilisation. For terrestrial (land) animals internal fertilisation is clearly essential because sperm must reach eggs by swimming through the female reproductive tract in a fluid. Internal fertilisation increases the chances of gametes meeting and fusing successfully, so fewer gametes are needed. This therefore reduces the total cost of gamete production. Internal fertilisation also offers increased protection for gametes (from predation and abiotic threats).

Internal Fertilisation
Where there is internal fertilisation, females have the option of laying eggs from which the young latter hatch, or allowing embryonic development to occur within the reproductive tract before giving birth to well developed young.

Sharks have internal fertilisation with some species laying eggs, while others are live bearers.

Every species of bird is an egg layer.

In mammals the majority give birth to live young with a few laying eggs (e.g. duck-billed platypus and spiny anteater).

2. Courtship & Display

Sexual selection
Monograph page 42

Darwin recognised that any characteristic which increases the breeding success of one gender will tend to become more common in successive generations of that gender. In 1871 Charles Darwin coined the term sexual selection to describe the evolution of different adaptations in the two genders that increase the likelihood of reproductive success.

Sexual selection
The adaptations which increase the likelihood of reproductive success in males may not be useful adaptations in females and vice versa. As a result, the sexes of many species differ in both appearance and behaviour. Differences between the two sexes of the same species is called sexual dimorphism (this is a product of sexual selection).

Rainforest frogs mating

Leaf mimic bush crickets feeding on a leaf

Sexual selection: sexual dimorphism

Generally females tend to be inconspicuous.
What is the advantage of this?

This ensures the female is camouflaged.

Camouflage decreases chances of predation and improves the

survival chance for the female and her young.

http://www.bbc.co.uk/nature/life/ Western_Capercaillie#p0085bs3

Courtship and display

Monograph p42-47

Given the costs and potential benefits associated with reproduction, sexual selection has resulted in some complex and lengthy courtship behaviours involving very elaborate displays. These are usually shown by males towards females.

Courtship and display

Sexual selection can take several forms:
1. Male-male rivalry
Competition between males for females; this occurs when males can control access to females through dominance. Through fighting males can increase their access to females.

Selection for fighting ability results in males that are often much larger than females.
The successful males will be those that are stronger or have greater weaponry, and these are fitness characteristics that will be advantageous to offspring. In some species competition is so intense that it frequently results in injury and sometimes death.

Male-male rivalry: Weapons size and injury

Gladiator frog, red deer and elephant seals are species in which males compete intensely with one another to achieve matings with females.

This has led to fighting adaptations in the three species in the form of large size, weapons or both.

Male-male rivalry: Weapons size and injury

Gladiator frogs are so called because males have a sharp bony spine on each hand, which is used as a weapon in fights between males over females and nests.
Extremely fierce fights over females often result in males being wounded or killed.

Male-male rivalry: Weapons size and injury

Red deer males are larger than females and bear antlers during the breeding season (rut). Nearly a quarter of mature stags may suffer injuries, including a broken antler, blinding or a broken leg.
Red deer are the largest of the UK's resident deer species. Their impressive autumn rut is a top British wildlife spectacle, particularly familiar in Scotland where red deer are widespread and common. During the rut, mature stags invest much time and effort into competing with other males for access to females. Roaring contests and parallel walking allow males to size each other up without violence. However, conflict can soon escalate between evenly matched stags leading to serious injuries and even death.

Male-male rivalry results in the sexually dimorphic evolution of weaponry in male red deer . The females lack antlers and they have a smaller body size.

Male-male rivalry: Weapons size and injury

Elephant seal males are much larger than females. During the breeding season most males show fresh wounds on their necks inflected by the bites of rival males. The wounds can be severe.

http://www.bbc.co.uk/nature/li fe/Southern_Elephant_Seal# p00l714p

Male-male rivalry: Sperm competition

Evidence has shown that intense competition between males may extend into the females reproductive tract where the sperm of rival males are in competition for access to eggs. This has been termed sperm competition and is the result of a time delay which may be of days or even weeks between mating and egg fertilisation. This gives the opportunity for sperm of later matings to supplant those already present. For example in some species of damselfly, males have special devices for removing the sperm of rival males from the female reproductive tract before introducing their own.

Courtship and display

2. Female choice of males
Having invested heavily in their eggs females could waste that investment by mating with a poor quality male.

Female choice occurs when males cannot control access to females.

In this situation females may choose males on the basis of the males fitness or his resources (i.e. those judged to be of high quality). This is sexual selection by female choice and leads to males having more conspicuous markings, structures and behaviours in order to attract females.
Bird of paradise display Tragopan display

Female choice in peafowl

Monograph Page 46-47

Read about good genes and female choice in peafowl. Study the graphs. Write brief summary notes.

3. Avoidance of inbreeding

Avoidance of inbreeding
Monograph page 47-51

Inbreeding is the sexual reproduction of close relatives e.g. brothers and sisters (or of individuals with very similar genomes). Inbreeding tends to lead to highly homozygous genomes and reduces the genetic variability in a population. Inbreeding increases the expression of disadvantageous or lethal recessive genes. This results in lower fitness and reduced breeding success in individuals who are homozygous for these alleles. Evidence of this comes from studies of inbred populations of lions and great tits (see monograph page 48).

Inbreed populations maintain high levels of homozygous alleles.

Outbreed populations maintain high levels of heterozygous alleles.

Inbreeding Depression

This lowering of fitness is called inbreeding depression. We should expect natural selection to favour the evolution of behaviour that reduces high levels of inbreeding. However, there may also be disadvantages to individuals that show extreme tendencies to outbreed. When an animal population has been living in the same environment for many generations, individuals will have become adapted to it in a variety of ways through natural selection. In this situation a strong tendency towards outbreeding would disrupt this collection of adaptations, leading to outbreeding depression. Evidence to support this comes from female white-crowed sparrows. These birds are more attracted to the songs of males of their local area than those of males from further away. This has led to the suggestion that selection might favour an optimal level of outbreeding (see monograph page 48-49).

Avoidance of inbreeding
Most species have mechanisms to ensure they mate within the same species but avoid mating with related individuals (i.e. avoid inbreeding). These mechanisms may involve dispersal of one sex only.

Avoidance of Inbreeding: Natal dispersal

Monograph Page 50-51

Read about natal dispersal in your monograph. Study the graph. Write brief summary notes. Include in your notes the difference between polygamy and monogamy.

Your notes should include the following points: Avoidance of inbreeding: Natal Dispersal
Natal dispersal = close family members disperse from where they are born (natal home) before they try to breed. Both sexes should benefit but studies of birds and mammals shows there are interesting differences between the sexes due to the relative costs and benefits of dispersal. In mammals males are found to be the sex which tends to disperse. In mammals the common mating system is polygamy males mating with several females in one season. As a consequence young males will need to compete with older males in their natal group. It would therefore be beneficial to a young male to disperse. Females are not in competition for mates because males can mate with several females, in addition since males disperse females do not risk inbreeding by staying where they are.

Example: Male natal dispersal in Lions

As cubs grow up in the pride they show the typical mammal pattern of dispersal.

Avoidance of inbreeding continued...

In bird species - female dispersal is the most common. In birds the common mating system is monogamy males have only one female mate forming a breeding pair. Occupying a breeding territory. The male usually has to acquire a territory in order to attract a mate. It has been suggested that for male birds, knowledge of the local area or ability to inherit part of a fathers territory could be an important reason for not dispersing. As a consequence females show natal dispersal to avoid inbreeding.

Test your knowledge...

1. Using a named example of an animal species, describe a characteristic of males used by females to choose their mate. 2. What term is used to describe the condition where there are distinct differences between males and females of the same species? 3. State one function of male displays. 4. How do females benefit from being inconspicuous? 5. Females often provide shelter, food and protection for their young. Describe another way in which female investment is likely to be greater than that of the male parent. 6. Male-male rivalry results in the evolution of weaponry in males. Give an example of this in a named animal. 7. Give two reasons why it is important that animals avoid inbreeding. 8. Distinguish between the mating systems monogamy and polygamy.

Past paper practice Discuss the role of dispersal in the avoidance of inbreeding in birds and mammals. Why is it important for animals to avoid inbreeding and how is this achieved? (4 marks) 1. High levels of homozygosity in inbred populations 2. Heterozygosity in outbred populations 3. Inbreeding increases expression of disadvantageous/ lethal recessive genes OR converse for outbreeding 4. Inbreeding results in lower fitness/inbreeding depression/reduced breeding success OR converse for outbreeding 5. Male dispersal in mammals/or example 6. Female dispersal in birds/or example

BBC - Wildlife Finder - Parthenogenetic aphids BBC - Wildlife Finder - Parthenogenetic lizards
BBC - Wildlife Finder - Coral spawning BBC - Wildlife Finder - Courtship display BBC - Wildlife Finder - Capercaillie lecking BBC - Wildlife Finder - Bower bird