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1 Communication Chapter 12 Photo Copyright R. Hays Cummins.

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2 1 Communication Chapter 12 Photo Copyright R. Hays Cummins

3 2 Communication What is communication? What is communication? Types (traits vs. behaviours), classification Types (traits vs. behaviours), classification How signals convey meaning How signals convey meaning Channels of communication (methods) Channels of communication (methods) Functions of communication (purposes) Functions of communication (purposes) Complex Communication (bee dance) Complex Communication (bee dance)

4 3 Definition Types of Communication States/Traits Behaviours (‘signals’) What is Communication? How is Communication Classified? Categories of Interaction (Wiley) Other views

5 4 What Is Communication? In biology, defined as: In biology, defined as: The transfer of information from one molecule, cell, or organism to another, as by chemical or electrical signals or by behaviours. (www.dictionary.com) The transfer of information from one molecule, cell, or organism to another, as by chemical or electrical signals or by behaviours. (www.dictionary.com) All communication involves a “sender” and a “receiver” All communication involves a “sender” and a “receiver”

6 5 Biological Communication Signal: The physical form in which a message is coded; any behaviour pattern that communicates something Discrete vs. Graded Discrete vs. Graded Discrete: “All or none” (e.g., FAPs) Discrete: “All or none” (e.g., FAPs) Graded: Varying in intensity and/or frequency; provides info about the strength of stimulus (e.g., Betta splendens display) Graded: Varying in intensity and/or frequency; provides info about the strength of stimulus (e.g., Betta splendens display)

7 6 Increasing intensity Discrete signal (ear position) Graded signal (mouth)

8 7 Biological Communication Displays are one type of signal Display: Any behaviour pattern especially adapted (i.e., modified through evolution) in physical form or frequency to function as a social signal in communication Display: Any behaviour pattern especially adapted (i.e., modified through evolution) in physical form or frequency to function as a social signal in communication e.g., spruce grouse males beating wings on air e.g., male dominance mounting in dogs

9 8 Two male Betta splendens showing fin flare and operculum (gill) flare Communicates willingness to fight; allows assessment of opponent without combat/injury

10 9 Wiley ( 1983 ): Categories of Interactions Terms used to describe interactions, depending on the value of the info to sender and receiver Terms used to describe interactions, depending on the value of the info to sender and receiver Value of info to SENDER: positive zero (negative) positive “True communication” “Manipulation” (“deceit”) zero (negative) “Eavesdropping” (“exploitation”) “Ignoring” (“spite”) Value of info to RECEIVER +/++/++/++/+ +/0 +/- 0/0 -/- 0/+-/+0/+-/+0/+-/+0/+-/+

11 10 Biological Communication How to classify/define communication? Value of information to each participant Value of information to each participant Cost & Benefit Analysis Both benefit = true communication; Both benefit = true communication; male bird song during mating season Sender benefits, receiver is unaffected or harmed = deceit or manipulation; e.g., killdeer performing “broken wing display” Sender benefits, receiver is unaffected or harmed = deceit or manipulation; e.g., killdeer performing “broken wing display”

12 11 Biological Communication Deceit Deceit Interspecific deceit more common Interspecific deceit more common predator-prey relationships predator-prey relationships Aquatic predators that attract prey using lures that look like insects Aquatic predators that attract prey using lures that look like insects Intraspecific “deceit” unlikely to spread in a population, except… Intraspecific “deceit” unlikely to spread in a population, except… ® Disney, Finding Nemo

13 12 Categories of Interactions Sender benefits, receiver is harmed (+/-): Sender benefits, receiver is harmed (+/-): Deceit Deceit Another example: Another example: “femme fatale” fireflies – females pretend to be receptive, attract and consume males of another species “femme fatale” fireflies – females pretend to be receptive, attract and consume males of another species

14 13 Biological Communication Value of information to each? Cont’d Sender unaffected or harmed, receiver benefits = exploitation or eavesdropping; Sender unaffected or harmed, receiver benefits = exploitation or eavesdropping; mouse rustles grass as running, allowing owl to hear it, catch it & eat it mouse rustles grass as running, allowing owl to hear it, catch it & eat it Neither benefit, both may be harmed = ignoring or spite; Neither benefit, both may be harmed = ignoring or spite; a bird ignores the cryptic colouration of a moth and tries to eat it a bird ignores the cryptic colouration of a moth and tries to eat it Monarch Butterfly Monarch Butterfly

15 14 Biological Communication Other ways to view communication: Other ways to view communication: Communication need not benefit both sender and receiver since natural selection operates mainly at the level of the individual (Dawkins & Krebs); Communication need not benefit both sender and receiver since natural selection operates mainly at the level of the individual (Dawkins & Krebs); Displays evolve to “waste” a minimum of energy of sender… persuade the receiver, not “inform” (exaggeration and redundancy of signals); Displays evolve to “waste” a minimum of energy of sender… persuade the receiver, not “inform” (exaggeration and redundancy of signals); roar of male lion roar of male lion

16 15 Other ways to view communication: Other ways to view communication: Displays – have evolved to maximize the effectiveness of information transfer, to the benefit of both sender and receiver Displays – have evolved to maximize the effectiveness of information transfer, to the benefit of both sender and receiver coevolution of signals; each species acts as a strong selection force on the other; coevolution of signals; each species acts as a strong selection force on the other; Problem: this definition cannot encompass “lying” Problem: this definition cannot encompass “lying” Biological Communication

17 16 How Signals Convey Meaning Distance & Duration (signal type) Distance & Duration (signal type) Can travel far (e.g. scent), or operate over short distances (e.g., visual signals) Can travel far (e.g. scent), or operate over short distances (e.g., visual signals) Can be long-lasting (e.g., pheromones entire breeding season) vs. short-term (e.g., calls/ vocalizations) Can be long-lasting (e.g., pheromones entire breeding season) vs. short-term (e.g., calls/ vocalizations)

18 17 How Signals Convey Meaning Individuals can increase information content of signals by… combining 2+ signals = composite signal Individuals can increase information content of signals by… combining 2+ signals = composite signal Zebra: meaning of “open mouth” signal depends on ear position (forward = friendly; backwards = hostile) Zebra: meaning of “open mouth” signal depends on ear position (forward = friendly; backwards = hostile) …changing syntax, or sequence of displays …changing syntax, or sequence of displays A before B is NOT = B before A A before B is NOT = B before A

19 18 How Signals Convey Meaning Information in a signal can be context- dependent; Information in a signal can be context- dependent; E.g., “lion’s roar” : aggressive, maintaining contact with pride members, or indication to neighbours to keep a good distance E.g., “lion’s roar” : aggressive, maintaining contact with pride members, or indication to neighbours to keep a good distance

20 19 Types of Communication (How can a message be sent?) States or Traits: States or Traits: Sign stimuli – Ch.’s 2 & 10 Sign stimuli – Ch.’s 2 & 10 Aposematism & Batesian mimicry – Ch. 15 Aposematism & Batesian mimicry – Ch. 15 Indicator traits – Ch. 17 (to come) Indicator traits – Ch. 17 (to come) Behaviours: (i.e., “does”) Behaviours: (i.e., “does”) Signals Signals Some FAPs Some FAPs Displays Displays

21 20 Three-spined stickleback (Ch. 2, p. 15) Three-spined stickleback (Ch. 2, p. 15) Enlarged belly of female = Sign stimulus (honest sign that female is ready to spawn) Enlarged belly of female = Sign stimulus (honest sign that female is ready to spawn) Sign Stimuli

22 21 Sign Stimuli Not a behaviour, a state (e.g., enlarged belly) Not a behaviour, a state (e.g., enlarged belly) May provide info about physical state of bearer May provide info about physical state of bearer NOTE: Not all sign stimuli convey information, and not all FAPs act as signals NOTE: Not all sign stimuli convey information, and not all FAPs act as signals E.g., egg retrieval in gulls E.g., egg retrieval in gulls If info is exchanged between conspecifics when sign stimulus or FAP occurs = communication If info is exchanged between conspecifics when sign stimulus or FAP occurs = communication

23 22 Aposematism (warning colouration) Conspicuous markings of noxious animals are easily recognizable and avoidable by predators Conspicuous markings of noxious animals are easily recognizable and avoidable by predators Convey information about unpalatability Convey information about unpalatability Batesian mimicry = manipulation/deceit (lying) Batesian mimicry = manipulation/deceit (lying) pinker.wjh.harvard.edu/ photos/santa_barbara_c...

24 23 Communication What is communication? What is communication? Types (traits vs. behaviours), classification Types (traits vs. behaviours), classification How signals convey meaning How signals convey meaning Channels of communication (methods) Channels of communication (methods) Functions of communication (purposes) Functions of communication (purposes) Complex Communication (bee dance) Complex Communication (bee dance)

25 24 Channels of Communication Odour Odour Sound (ultrasonic, sonic, and infrasonic) Sound (ultrasonic, sonic, and infrasonic) Seismic vibrations Seismic vibrations Touch Touch Electric field Electric field Visual Visual

26 25 Properties of Sensory Channels Range (distance covered) – short to long Range (distance covered) – short to long Transmission rate (how long does it take to get from sender to receiver?) – slow to fast Transmission rate (how long does it take to get from sender to receiver?) – slow to fast Can it travel around objects? – yes or no Can it travel around objects? – yes or no Can it be used at night? (yes, no, or sometimes) Can it be used at night? (yes, no, or sometimes) Fade-out time (how long does it last? – slow to fast) Fade-out time (how long does it last? – slow to fast) Locate sender? (easy to difficult) Locate sender? (easy to difficult) Cost to send signal (low to high) Cost to send signal (low to high) Complexity of message (low to high) Complexity of message (low to high)

27 26 Audition: Properties Auditory communication Auditory communication Range? Range? long long Travels around objects? Travels around objects? Yes Yes Useful at night? Useful at night? Yes Yes Fade-out time? Fade-out time? Fast Fast Cost to sender? Cost to sender? High High

28 27 Odour Used by most species (evolutionarily “old”) Used by most species (evolutionarily “old”) Usually involves pheromones: Usually involves pheromones: a species-specific odour cue released by animals that influences the behaviour and/or physiology of conspecifics a species-specific odour cue released by animals that influences the behaviour and/or physiology of conspecifics

29 28 Odour Functions: Functions: Group spacing – marking territories Group spacing – marking territories Recognition – identifying conspecifics, kin, individuals Recognition – identifying conspecifics, kin, individuals Reproduction – attracting mates, altering physiology (e.g., levels of sex hormones) Reproduction – attracting mates, altering physiology (e.g., levels of sex hormones) Alarm – fear pheromone (e.g., inverts) Alarm – fear pheromone (e.g., inverts)

30 29 Odour Pheromones often present in urine of mammals (& anal glands) Pheromones often present in urine of mammals (& anal glands) Two types: Two types: priming (generalized response) priming (generalized response) signaling (immediate specific response) signaling (immediate specific response) “Flehmen” response (vomeronasal organ) “Flehmen” response (vomeronasal organ)

31 30 Odour: Properties long range long range slow transmission rate slow transmission rate travels around objects travels around objects useful at night useful at night slow fade-out time slow fade-out time low cost to sender low cost to sender unable to locate sender unable to locate sender simple message only simple message only

32 31 Sound Can occur within or outside of human “umwelt” (~20-20,000 Hz) Can occur within or outside of human “umwelt” (~20-20,000 Hz) ultrasonic (high frequency - e.g., echolocation) ultrasonic (high frequency - e.g., echolocation) infrasonic (low frequency - e.g., African elephant rumbles, whale song) infrasonic (low frequency - e.g., African elephant rumbles, whale song)

33 32 Sound Can be produced by: Can be produced by: a single organ ( vocalizations) a single organ ( vocalizations) stridulation (e.g., cricket “chirp”) stridulation (e.g., cricket “chirp”) other means (e.g., drumming by Northern Flickers/ woodpeckers) other means (e.g., drumming by Northern Flickers/ woodpeckers)

34 33 Seismic Vibration Substrate-borne vibrations detected by animals Substrate-borne vibrations detected by animals White-lipped male frogs make thumps, females extremely sensitive to vibrations White-lipped male frogs make thumps, females extremely sensitive to vibrations Spiders catch prey by detecting vibrations Spiders catch prey by detecting vibrations Male orb-weaving spiders “pluck” at web to bring female out for mating Male orb-weaving spiders “pluck” at web to bring female out for mating

35 34 Sound/Vibration Properties long range long range fast transmission rate fast transmission rate travels around objects travels around objects useful at night useful at night fast fade-out time fast fade-out time high cost to sender high cost to sender may be able to locate sender may be able to locate sender sound = complex message, vibration = simple message sound = complex message, vibration = simple message

36 35 Touch Tactile communication (physical contact) Tactile communication (physical contact) Common in invertebrates: Antennae, tube feet, etc. (also pick up chemical cues) Common in invertebrates: Antennae, tube feet, etc. (also pick up chemical cues) Vertebrate copulation  tactile stimulation elicits lordosis response in female mammals Vertebrate copulation  tactile stimulation elicits lordosis response in female mammals Touch (e.g., vaginal stimulation) can induce ovulation in some species (induced ovulators) Touch (e.g., vaginal stimulation) can induce ovulation in some species (induced ovulators)

37 36 Touch: Properties short range short range fast transmission rate fast transmission rate can’t travel around objects can’t travel around objects useful at night useful at night fast fade-out time fast fade-out time low cost to sender low cost to sender easy to locate sender easy to locate sender Simple message only Simple message only

38 37 Visual Communication Markings, patterns, colour require vision Markings, patterns, colour require vision Mostly used by diurnal species (visual displays), e.g., aggressive displays of SFF Mostly used by diurnal species (visual displays), e.g., aggressive displays of SFF Exception: fireflies Exception: fireflies species-specific flashing to attract mates and during courtship species-specific flashing to attract mates and during courtship “femme fatales” of Photuris versicolor (deceit) “femme fatales” of Photuris versicolor (deceit)

39 38 Vision: Properties medium range medium range fast transmission rate fast transmission rate can’t travel around objects can’t travel around objects little use at night little use at night fast fade-out time fast fade-out time cost to sender medium cost to sender medium easy to locate sender easy to locate sender allows complex messages allows complex messages

40 39 Electric Field Some sharks and electric fish detect prey’s electric field using electroreceptors Some sharks and electric fish detect prey’s electric field using electroreceptors Strongly electric fish (e.g., electric eel) have EODs (electric organ discharges of 100’s V) Strongly electric fish (e.g., electric eel) have EODs (electric organ discharges of 100’s V) Social communication possible by altering wavelength/ pulse duration Social communication possible by altering wavelength/ pulse duration WhoZoo(http://whozoo.org)

41 40 Electric Field: Properties short range short range fast transmission rate fast transmission rate travels around (and through) objects travels around (and through) objects useful at night/in murky water useful at night/in murky water fast fade-out time fast fade-out time cost to sender varies cost to sender varies easy to locate sender easy to locate sender Simple message only Simple message only

42 41 Functions of Communication 1.Group spacing & coordination 2.Recognition 3.Reproduction (identifying/attracting mates) 4.Agonism and social status 5.Alarm (warning) 6.Finding food 7.Soliciting play 8.Giving & soliciting care 9.Synchronization of hatching

43 42 Functions of Communication Group Spacing & Coordination Group Spacing & Coordination Spacing signals of primates: Spacing signals of primates: Distance-increasing (branch shaking) Distance-increasing (branch shaking) Distance-maintaining (dawn chorus) Distance-maintaining (dawn chorus) Distance-reducing (contact/lost calls) Distance-reducing (contact/lost calls) Proximity-maintaining (social grooming) Proximity-maintaining (social grooming)

44 43 Functions of Communication Recognition Recognition Species Recognition Species Recognition Neighbour Recognition Neighbour Recognition Individual Recognition Individual Recognition Kin Recognition Kin Recognition

45 44 Functions of Communication Reproduction Reproduction Courting displays, copulation- allow assessments of species, physical condition, “quality” of mate Courting displays, copulation- allow assessments of species, physical condition, “quality” of mate

46 45 Indicator Traits ( Sexual Selection – Ch. 17 ) Provide info to members of the opposite sex about the physical condition of the bearer Provide info to members of the opposite sex about the physical condition of the bearer Choosier sex (usually females) show preference for most extravagant traits Choosier sex (usually females) show preference for most extravagant traits E.g., brightest colours, largest antlers, longest tail feathers, etc. E.g., brightest colours, largest antlers, longest tail feathers, etc.

47 46 Keeping signals HONEST: Zahavi & Zahavi’s “Handicap Principle” Zahavi & Zahavi’s “Handicap Principle” If signals are costly to produce (peacock tail), they tend to be honest indicators of condition or ability (i.e., in good health, well-nourished and free of parasites) (sexual selection)

48 47 Functions of Communication Social Status/ Agonism Social Status/ Agonism Dominance/submissive displays- avoid costly fighting Dominance/submissive displays- avoid costly fighting Traits may also indicate social status (bigger, stronger males perceived as dominant) e.g., antler size in moose Traits may also indicate social status (bigger, stronger males perceived as dominant) e.g., antler size in moose wolves and dogs wolves and dogs

49 48 Female Belding's ground squirrel (Spermophilus beldingi) emitting an alarm call. (Sherman) Alarm calls are given to warn other individuals about the presence of a nearby predator and are usually directed toward close kin.Alarm calls are given to warn other individuals about the presence of a nearby predator and are usually directed toward close kin. giving an alarm call increases the chance that the predator will attack the callergiving an alarm call increases the chance that the predator will attack the caller Functions of Communication Alarm Altruism or Selfishness?

50 49 Functions of Communication Alarm Alarm Vervet monkeys: Seyfarth & Cheney Vervet monkeys: Seyfarth & Cheney “semantics”- different calls for different predators (avian, ground) “semantics”- different calls for different predators (avian, ground) Pheromones as chemical alarm substances Pheromones as chemical alarm substances inverts; mammals inverts; mammals Sea urchins steer clear Sea urchins steer clear of an area where a urchin has just been crushed

51 50 Functions of Communication Finding Food Finding Food “Information Centre Hypothesis” “Information Centre Hypothesis” colonial organisms colonial organisms (e.g., communally roosting weaver (e.g., communally roosting weaver birds) hunting packs hunting packs (African wild dogs) (African wild dogs) Giving/Soliciting Care, Play; Giving/Soliciting Care, Play; Synchronization of Hatching (precocial birds – vocalize prior to hatching) Synchronization of Hatching (precocial birds – vocalize prior to hatching)

52 51 Evolution of Displays Communicative displays are believed to have evolved from noncommunicative behaviours through the process of ritualization: Communicative displays are believed to have evolved from noncommunicative behaviours through the process of ritualization: Ritualization = Evolutionary process by which behaviour patterns become modified to serve as communication signals. Ritualization = Evolutionary process by which behaviour patterns become modified to serve as communication signals.

53 52 Ritualization (example) Food calling behaviour still seen in roosters may be evolutionary origin of courtship display seen in peacocks Food calling behaviour still seen in roosters may be evolutionary origin of courtship display seen in peacocks i.e., peacocks’ ancestors may have performed food-calling behaviour, which became ritualized into bow/feather spread display presently used i.e., peacocks’ ancestors may have performed food-calling behaviour, which became ritualized into bow/feather spread display presently used

54 53 Evolution of Displays “freezing” of movement into postures “freezing” of movement into postures Stereotyping of behaviour (stays constant even under different levels of motivation) Stereotyping of behaviour (stays constant even under different levels of motivation) Development of conspicuous body structures- enlarged claws (fiddler crab), ornamental feathers (peacocks, male ducks), manes (lion), exaggerated fins (Betta) Development of conspicuous body structures- enlarged claws (fiddler crab), ornamental feathers (peacocks, male ducks), manes (lion), exaggerated fins (Betta) Crested Auklet (I. Jones)

55 54 Displays may have arisen from… Intention movements - e.g., “powk” calls of BL Kittiwakes -- gannets Intention movements - e.g., “powk” calls of BL Kittiwakes -- gannetsOR Displacement activities- innate, stereotypical responses to stimuli that seem inappropriate to the situation Displacement activities- innate, stereotypical responses to stimuli that seem inappropriate to the situation Some degree of conflict in organism’s motivation- e.g., strange female approaches: is a sexual or agonstic response needed? Some degree of conflict in organism’s motivation- e.g., strange female approaches: is a sexual or agonstic response needed?

56 55 Displacement Activities Evolved into courtship display: During courtship, males may preen or touch own feathers (sham preening) Evolved into courtship display: During courtship, males may preen or touch own feathers (sham preening) Conspicuous feathers (sailfeathers) may have evolved and are exposed during this behaviour Conspicuous feathers (sailfeathers) may have evolved and are exposed during this behaviour Mandarin Duck

57 56 Receiver bias - the increased sensitivity of perceptual systems to certain stimuli as a result of natural selection Receiver bias - the increased sensitivity of perceptual systems to certain stimuli as a result of natural selection  Such biases may influence the evolution of communicative displays  E.g., insect feeds mostly on yellow flowers, adapted to be sensitive to yellow in general, males with yellow are preferred, and may exploit this to attract females Other origins of displays:

58 57 Complex Communication

59 58 Honeybee communicative dances “Round Dance” Food nearby; less detailed “Waggle Dance” Food >200m away; more detailed

60 59 Photo Caption: Diagram of the honeybee dance. (Credit: P. Kirk Visscher.) Ceiling of hive = direction of sun (reference)  Angle of straight run (on vertical) = direction of food source relative to sun

61 60 direction and duration of the straight run is criticaldirection and duration of the straight run is critical antennae of hive members maintain contact with the dancer and taste samples of regurgitated foodantennae of hive members maintain contact with the dancer and taste samples of regurgitated food


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