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Postcopulatory Sexual Selection
Long Miao ([email protected])Institute of Biophysics, CAS
April 25, 2011
Male-Male Competition
• Andersson, 1994 “Sexual Selection”• 5 fundamental areas of male-male
competition – Scrambles– Endurance Rivalry– Contests– Sperm competition– Mate choice (Female choice)
Sperm Competition
• Competition between the sperm of 2 or more males within the reproductive tract of the female
• Typically Post copulatory sperm competition– Competition between males after copulation
• With few ova= huge battle– promiscuity go for broke
• Involves– Physiological events that occur after
insemination– Behaviors before & after copulation– Morphology
The Father of Sperm Competition
• Geoff Parker, 1970• Studied sexual selection in Yellow
Dungfly• Looked at patterns of male-male
competition
Yellow Dungfly
What’s poo got to do with it…
• Female arrives at dung patty ready to mate
• After copulation lays eggs on dung• Males mount & copulate with female as
soon as she arrives• Larger males removes male from female &
copulates with her• Females mate with multiple males
throughout day
Cow Patties
Yellow Dungfly
Parker’s Observations• Under such extreme male male
competition Parker realized competition between males continues through action of their sperm
• Copulation is not enough, sperm must make it to the egg
• Defined sperm competition– The competition between ejaculates of
different males for the fertilization of a female’s eggs
Parker’s Observations
• Sexual selection occurs in the reproductive tract– Fertilization not copulation is the driving force– Males “win” by
• Inseminating more sperm per ejaculate• Faster swimming sperm• Inseminate moderate amounts of sperm & remove
sperm from the last guy• Disabling rival sperm
• Sperm competition results in opposing selection forces in males
The Trade-off• Selection favors BOTH
– Males that fertilize females that have been inseminated by other males
– Males who prevent females they have inseminated from being fertilized by another male
• Results in counter adaptations among competing male strategies– Ex producing larger ejaculates vs better mate guarding
strategies• What?
– Males want to go around & fertilize as many females as possible, but this is hard to do while they are mate guarding
Darwin & Sperm Competition
2 General categories of Sperm Competition
• Adaptations that preventing rival sperm from copulating with the mate– Mate guarding, sequestering, frequent
copulation, production of mating plug, displacing rival sperm
• Adaptations that increase the Ability of one male’s sperm to outcompete the sperm of another male– production of abundant sperm, fast sperm
Cuckoldry
• Female infidelity• Derived from
cuckoo• Male loses some of
his reproductive fitness to another male
Sperm Precedence
• The sperm that fertilizes the egg effected by the order of fertilization
Sperm Precedence
• First male precedence– First male fertilizes the majority of eggs
• Last male precedence– Last male fertilizes the majority of eggs
• promiscuity
Sperm Competition
• Adaptations that preventing rival males from copulating with the mate– Mate Guarding
Mate Guarding
• Male increases the amount of time he spends in close association with a fertile female
• Male guards female partners from mating with other males
• Tactic for reducing or preventing sperm competition
• Typically occurs when female is receptive for a short time period
• Costs of mate guarding– Interferes with male foraging– Vulnerable to predators
Sand Martin
Sand Martin
• Male guards female 7-8 days• Begins few days before she lays first
egg• Male remains next to female during
her fertile period• Male presence reduces risk other
males copulating with female
Red-winged Blackbird
• David Westneat• Male guards females
– Male experimentally removed from territory
– Female copulations with other males increased 100X
• Males Compromise guarding for foraging time– When males given food
paternity went up
Mate guarding strategies
• Mate grasping• Prolonged
copulations• Frequent
Copulations• Mate sequestering• Mating plugs
Yellow DungflyMate Grasping
Male remains on female’s back as she lays eggs
Pre-copulatory Guarding
• Occurs in species where the first male has the best chance fertilizing the female’s egg
Freshwater Shrimp
• Males use specialized legs to carry females around
• Males attach to female 9 days before she is receptive
Mate Guarding- Amplexus
• Salamanders & Frogs
Mate Guarding- Prolonged Copulation
• Male & females remain in copulation • May last hours to days• Male prevents other males by
staying attached
Yellow Dungfly
Prolonged copulation…male remains with female 40min
Prolonged Copulation
Frequent Copulations
• Male who increases copulation events with female prevents mating with other males
• Animals frequently mate more often than necessary for fertilization
Mate Guarding- Frequent Copulation
• Giant water bug aka “toe biter”• Male fertilizes females eggs• As female lays eggs she sticks them
to the males back– Lays 1-2 eggs at a time
• Cycle can last up to 36 hours (144 eggs)
• Male fertilizes over 99% of eggs
Giant Water Bug
Days….
Frequent Copulations
• Mountain Lion• Ave 9 copulations
per hour
Mate Sequestering
• Male limits female contact with other males by isolating her– Physical isolation– Chemical isolation
Mate Sequestering Physical Isolation
Mate Sequestering Physical Isolation
• Douglas Fir Bark Beetle• Male sequesters female in gallery
Mate Sequestering Chemical isolation
• Reducing Attractiveness of mates• “Anti-aphrodisiac”• Substances that reduce the females
attractiveness• Ex. Make her smell like a male• Reduce her pheromonal output
Chemical Mate Guarding
• Green veined white butterfly• Males inject methyl salicylate along
with sperm– Chemical compound derived from host
plant
• Smell repels other males
Green veined white butterfly
Spider- Linyphia litigiosa
• Male disrupts female’s chemical signals
• Male that finds virgin female will reduce evaporation of attractive pheromones from her web by packing it into a tight mass
Central American Bee
• Mandibular gland rubs secretions onto female makes her smell like a male
• Conceals female
Darkling Beetle
• Male rubs anti-aphrodisiac pheromone with genitals
• Makes female less attractive to other males
Mating Plugs
• Aka copulatory plugs• Male produced substances or
structures which are transferred to the female after copulation
• Seal genital opening• Ex guinea pig, chimpanzee, lemur,
masked palm civet,
Red-sided Garter Snake
Red-sided Garter Snake
• Intense male competition• Male produces gelatinous secretion
in ejaculate• Secretion hardens in the female
oviduct
Ground Squirrels
• Intense competition for females• Male emerges from hibernation first• Testes grow• Females emerge & within 4 days are
receptive– 1-2 hours
Mating Plugs
Mating Plugs
Orb Spider Argiope bruennichi
Orb Spider Argiope bruennichi
• Male transfers sperm packet to female reproductive tract using pedipalp
• Pedipalp breaks off in female & created mating plug
Orb Spider Argiope bruennichi
Honey Bee
• Uses own body as a copulatory plug
• Male dies after reproduction
• Harvester ant
Fly with no common name
• Ceratopogonid Johannseniella nitida
• Tiny male crawls into female genital opening to create mating plug
Ring-tailed Lemur
Other Functions of Mating Plugs…
• Sperm storage reservoir• Reduce leakage
Displace Rival Sperm
• displace a previous male’s sperm• Mechanism for last male precedence
– Last male fertilizes the majority of the offspring by removing previos males’ sperm
• Adaptations– Physical– Chemical
Black-winged Damselfly
• Females mate with multiple males within a few hours
• Sperm storage in spermotheca• Male penis shaped like scrub brush• During copulation male abdomen pumps
up & down “scrubbing” out the sperm of other males– 90-100% of sperm removed
What’s a Spermatheca?
Black-winged Damselfly
Fertilization success depends on:
1. Physical traits of sperm: numbers of sperm inseminated,
quality, size, morphology, speed, cooperation of close-related
2. Seminal fluid and its components produced by several accessory glands in male body
3. Cryptic female choice