Harder, better, faster, stronger: why some crickets put more effort into attracting females

Body morphology, energy stores, and muscle enzyme activity explain cricket acoustic mate attraction signalling variation

A new article in PLOS ONE written by

Ian Thomson, Charles Darveau, and Sue Bertram

Click here for a copy of our PLOS ONE article

Click here for the Bertram Lab website

To get lucky, males crickets have to chirp harder, better, faster, and stronger.

Figure 1: To get lucky, males crickets have to chirp harder, better, faster, and stronger.

In nature, females choose males for many reasons. Chosen males may have bright plumage, they may be good fighters, or they may successfully serenade females with their song. Male field crickets try to convince females that they are the best mate by producing more superior chirps than other males. Females tend to prefer males whose signals are harder, better, faster, and stronger (or more accurately longer in duration, more frequent throughout the night, at higher pulse rates, and louder; Figure 1).

Since male crickets with the best signals typically mate most often, we should expect that all males should signal with high effort. But instead, males exhibit substantial variation in how often they signal, with some males signaling for much of their adult life while others rarely signal. Why does variation in cricket signalling behaviour persist?

One appealing solution to this problem is the genic capture hypothesis. Genic capture assumes that variation in sexually selected traits is constrained by an organism’s condition. Instead of genes directly determining a male’s ability to signal for a mate, the genes code for a whole bunch of different factors that affect a cricket’s condition (Figure 2).

condition

Figure 2: Traits are dependent on underlying physiological, biochemical, behavioural, morphological traits that all combine to make up an organism’s “condition

These factors could be physiological, biochemical (one cricket may have more active enzymes than another), or morphological (one cricket may have more fat/ carbohydrate stored than another) in nature. So instead of genes directly determining how a male signals, variation in condition may drive variation in signaling effort.

composition

Figure 3: Crickets were dissected, and assays measured the total amounts of carbohydrates, lipids, and glycogen.

To measure male crickets’ condition (and by doing so address the proximate causes underlying variation in signalling effort) we quantified the total energy stores in the abdomen and thorax (Figure 3) and the maximal activities of key metabolic enzymes in the calling muscles in two species of field cricket and assessed whether they correlated with signalling variation. We did this work on chirping male Jamaican field crickets (Gryllus assimilis) and trilling male Texas field crickets (Gryllus texensis) (Figure 4).

Figure 4: The chirping G. assimilis and the trilling G. texensis were used in this study

We hypothesized that

1) There will be a correlation between variation in cricket sexual signalling quantity (time spent signalling in a given night) and quality (how attractive females find the signalling) and variation in signalling muscle enzymes

2) Variation in sexual signalling quantity and quality would be correlated with variation in capacity to accumulate and store carbohydrates and/or lipids.

We found that chirping G. assimilis primarily fuelled signalling with carbohydrate metabolism: individuals with increased thoracic glycogen stores signalled for mates with greater effort; individuals with greater glycogen phosphorylase (an enzyme used for carbohydrate metabolism) activity produced more attractive mating signals.

Conversely, the more energetic trilling G. texensis fuelled signalling with both lipid and carbohydrate metabolism: individuals with increased β-hydroxyacyl-CoA dehydrogenase (an enzyme used for lipid metabolism) activity and increased thoracic free carbohydrate content signalled for mates with greater effort; individuals with higher thoracic and abdominal carbohydrate content and higher abdominal lipid stores produced more attractive signals.

Our findings suggest variation in male reproductive success may be driven by hidden physiological trade-offs that affect the ability to uptake, retain, and use essential nutrients. Our findings reveal that a physiological perspective can help us to understand some of the causes of variation in behaviour.

ResearchBlogging.org

Ian R. Thomson, Charles A. Darveau, & Susan M Bertram (2014). Body Morphology, Energy Stores, and Muscle Enzyme Activity Explain Cricket Acoustic Mate Attraction Signaling Variation PLOS One : DOI: 10.1371/journal.pone.0090409

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Breaking up the Edge to Hide in Plain Sight

Image

A guest post by Rich Webster

Whether hiding from Lions on the Serengeti or dodging bullets on the battlefield, avoiding unwanted eyes should be high up on your priorities!

In nature, animals have evolved different strategies to achieve concealment. Resembling the appearance of ones surroundings (background matching) can reduce the chance of being detected. Surprisingly background matching is not a golden bullet solution to camouflage. Even species with faithful background matching can suffer from having visible edges, due to a small mismatch between where the animal starts and the background ends. For instance, predators can use these visible edges as cue to recognise camouflage prey from there characteristic shape.

Alternatively, an animal can trick the eye-of-the-beholder into seeing but failing-to-recognise its prey by masking distinctive features using disruptive coloration. A 100-year old theory speculated that background matching camouflage can benefit from a complimentary camouflage strategy of disrupting outlines visibility (Fig. 1).

Do edge patches contribute to camouflage through background matching or disruptive coloration? (a) White-tailed Ptarmigan Lagopus leucura, (b) Marai giraffe Giraffa camelopardalis tippelskirchi, (c) Canadian Disruptive Pattern (CADPAT), the Worlds first digital camouflage, (d) Emu chicks Dromaius noaehollandiae, (e) tree frog  Hyla versicolor. Photographs by Michael Webster (a-d) and Micheal Runtz (e).

Fig. 1. Do edge patches contribute to camouflage through background matching or disruptive coloration? (a) White-tailed Ptarmigan Lagopus leucura, (b) Marai giraffe Giraffa camelopardalis tippelskirchi, (c) Canadian Disruptive Pattern (CADPAT), the Worlds first digital camouflage, (d) Emu chicks Dromaius noaehollandiae, (e) tree frog Hyla versicolor. Photographs by Michael Webster (a-d) and Micheal Runtz (e).

Disruptive edge markings intersect animals’ outline, breakup its edge, making boundary and overall shape less recognisable. Whilst many animals, such as zebras, tigers and cuttlefish, have been proposed to have disruptive camouflage—not to mention a host of military uniforms and equipment—to date there is no evidence show that disruptive patterns are harder to recognise, a key prediction of this concept.

Here at Carleton University, Ottawa, Canada, we set out to test experimentally if disruptive camouflage truly misleads humans when searching for animals. Using humans hunting on computer screens (Fig. 2a) we looked at the survivorship of artificial moth targets with varying numbers of edge patches. Further, we used eye tracking technology to measure recognisability (Fig. 2b). Targets that were looked at for longer were assumed to be harder to recognise.

Figure 2 a) Photograph of a human subject hunting on a computer screen for an moth hidden (see grey arrow) on the tree image, b) a close up of the eye-tracking apparatus and the screen with an overlay showing where the subject was looking during their search.

Fig. 2. a) Photograph of a human subject hunting on a computer screen for an moth hidden (see grey arrow) on the tree image, b) a close up of the eye-tracking apparatus and the screen with an overlay showing where the subject was looking during their search.

We predicted that if number of edge patches improves camouflage by disruptive coloration, then targets with more edge intersecting patches should have a high survivorship due to impaired recognition.

Indeed, we found that targets with more edge intersecting patches taking longer to be found. This cannot be explained due to background matching alone because this was even the case for edge markings that were dissimilar from the background. Crucially, targets with more edge patches took longer to be recognised and were overlooked more often.

In our recent Biology Letters publication, we offer new evidence to support the hypothesis that disruptive coloration can achieve camouflage by masking animals’ outline visibility, which makes animals less recognisable.

Further, these novel methods provide a means to test if seemingly ‘disruptive’ markings, such as those of zebras, tigers or even soldiers uniforms function to disrupt recognition (Fig. 3).

Figure 3: Tiger stripes have long been speculated to break up the cats like shape. The next step that scientist studying camouflage will take is to apply their new methods of assessing camouflage, to start studying animals in the wild.

Fig. 3. Tiger stripes have long been speculated to break up the cats like shape. The next step that scientist studying camouflage will take is to apply their new methods of assessing camouflage, to start studying animals in the wild.

This research will enable us to better understand—and  design—camouflage patterns, as well as appreciate the beauty of animals colorations, that we sometimes don’t always see.

Listen to Rich Webster discuss this research in a 3 min YouTube video:

 

Richard Websterrichard.j.webster@gmail.com

Skype username: Richard.j.webster

Dr. Chris HassallC.Hassall@leeds.ac.uk

www.christopherhassall.com

Middle age females are most interested in mating: Lessons from a cricket

How age influences phonotaxis in virgin female Jamaican field crickets (Gryllus assimilis)

A new article in PeerJ written by Karen Pacheco, Jeff Dawson, Mike Jutting, and Sue Bertram

Click here for a copy of our PeerJ article

Click here for the Bertram Lab website

Female mate choice can heavily influence the evolution of male sexual signals. While mate choice was once thought to be a consistent and stable behaviour[1], research has shown mate selection is often very dynamic and variable throughout a female’s life. There are a number of environmental and internal factors that can affect a female’s mate choice (Figure 1). For instance, a female suddenly in danger of encountering a predator may alter her final mate decision[2]. A female’s past sexual experience[3] and her age[4] [5] [6] can also play a role in determining her mate choice. We currently have a vague understanding of how and why various intrinsic factors, such as female age, can cause variation in mate preferences and what the overall effect is on the evolutionary direction of male sexual traits.

Network of factors influencing choice

Figure 1. Network of some of the intrinsic and extrinsic factors that could influence the final mate choice expressed before copulation; the interaction of these mate preference cues can drive sexual selection[7].

In one of our studies, which was recently published in PeerJ, we examined how one of these factors, female age, influenced female mate preferences in the Jamaican field cricket, Gryllus assimilis. Research on other species including cockroaches (Nauphoeta cinerea)6 and guppies (Poecilia reticulate)5 has revealed that as females age, their health and likelihood of future reproduction progressively declines. Because of this, older females may be less choosy in selecting mates compared to younger females who have higher reproductive potential.

Trackball schematic

Figure 2. A) A cricket tethered on top of the sphere (part of trackball). An optical sensor picks up the movements made by the trackball, which is rotated by the cricket walking. This information is collected on a computer system[10].

Figure 2. B) Female cricket tethered on top of our trackball via spring attached to magnet.

Figure 2. B) Female cricket tethered on top of our trackball via spring attached to magnet.

In crickets and other acoustic insects, females find and assess the attractiveness of potential mates using acoustic signals produced by the males of their species (termed ‘phonotaxis’)[8] [9]. We tested a wide range of female ages and assessed female phonotactic response to a male acoustic mate attraction signal using a specially built apparatus called a trackball. The trackball enables us to suspend a female over rotatable sphere and measure her movement associated with her attempt to move towards a speaker playing a male’s acoustic signal (Figure 2). Using the trackball and associated computer software, we measured how fast a female moved towards the speaker (velocity), how far she moved (total path length), and whether she generally seemed to be attempting to move towards the speaker (net vector score).

We found that female age influenced phonotaxis response: older females (13 days post adult moult) moved further and at higher velocities than females in other age groups in an attempt to reach the speaker (Figure 3).

How age influences phonotaxis

Figure 3. How age affects female phonotaxis measures. Letters above each age reveal significant differences.

We also found that older females (10-13 days post adult moult) tended to more accurately orient themselves and move towards the active speaker than younger females (1-7 days post adult moult). The polar plots below (Figure 4) show the average direction and velocity (red arrows) towards the active speaker (0°) for females across all 10 age groups.

Polar Plots

Figure 4. Polar plots indicating speed and direction moved for each age group tested. Inner circular gridlines indicate velocity magnitude (cm/s) and tic marks around the outside of the circle represent velocity angle. Red arrows show the mean velocity for all 20 females in each age group. The large, multi-colored plot shows the differences in average velocity across age groups.

Overall, we found age influenced female G. assimilis phonotaxis, where middle aged adult females’ (10-13 days old) displayed the strongest phonotaxis compared to younger or older females. The high speed and long distance travelled observed in young females (1-7 day) may signify higher energy reserves9, but their random orientation suggests little interest in the standardized acoustic mate attraction signal. We provide two possible explanations for the young females’ inattention towards the standard mate attraction signal: stronger preferences and lack of maturity. Very young females may have greater requirements in mate selection and prefer more extreme male traits[11]. Alternatively, very young females may be too young to respond to the call because they are not reproductively mature yet. Future research should present young females with multiple male signals to determine if indeed they are reproductively immature or only phonolocating toward signals with extreme trait values. Conversely, the older females oriented toward the speaker, suggesting interest, but walked at reduced speeds and covered shorter distances, possibly indicating senescence[12]. Overall, our findings are consistent with most studies on how age influences mating preference: diminished choosiness with increasing age[13] [14]. Our study suggests 10- and 13-day females are most responsive when quantifying the preference landscape for G. assimilis sexual signals.

karen

Karen Pacheco is about to complete her MSc degree in Biology at Carleton University

ResearchBlogging.org
Karen Pacheco, Jeff Dawson, Mike Jutting, and Sue Bertram (2013). How age influences phonotaxis in virgin female Jamaican field crickets (Gryllus assimilis) PeerJ : 10.7717/peerj.130

[1] Andersson, M. 1994. Sexual Selection. Princeton: Princeton University Press.

[2] Magnhagen, C. 1991. Predation risk as a cost of reproduction. Trends in Ecology & Evolution, 6, 183-186.

[3] Dugatkin, L.A. 1992. Sexual selection and imitation: females copy the mate choice of others. The American Naturalist, 139, 1384-1389.

[4] Jouventin, P., LeQuette, B., & Dobson, F.S. 1999. Age-related mate choice in the wandering albatross. Animal Behaviour, 57, 1099-1106.

[5] Kodric-Brown, A., & Nicoletto, P.F. 2001. Age and experience affect female choice in the guppy (Poecilia reticulata). The American Naturalist, 157, 316-23.

[6] Moore, P.J., & Moore, A.J. 2001. Reproductive aging and mating: The ticking of the biological clock in female cockroaches. Proceedings of the National Academy of Sciences, 98, 9171-9176.

[7] Widemo, F., & Sæther, S. A. 1999. Beauty is in the eye of the beholder: causes and consequences of variation in mating preferences. Trends in Ecology and Evolution, 14, 26-31.

[8] Solymar, B., & Cade, W.H. 1990. Age of first mating in field crickets Gryllus integer (Orthoptern: Gryllidae). The Florida Entomologist, 73, 193-195.

[9] Prosser, M.R. 1994. Effect of Age On Female Choice in the Field Cricket, Gryllus integer. Unpublished Masters thesis, Brock University, St. Catharines, Ontario.

[10] Hedwig, B. & Poulet, J.F. 2004. Complex auditory behaviour emerges from simple reactive steering. Nature, 430, 781-785.

[11] Ryan, M.J., Keddy-Hector, A. 1992. Directional patterns of female mate choice and the role of sensory biases. The American Naturalist, 139, 4-35.

[12] Prosser, M.R., Murray, A.M., & Cade, W.H. 1997. The influence of female age on phonotaxis during single and multiple song presentations in the field cricket, Gryllus integer (Orthoptera: Gryllidae). Journal of Insect Behavior, 10, 437-449.

[13] Roff, D.A. 1992. The evolution of life histories: theory and analysis. London: Chapman & Hall.

[14] Mautz, B.S., & Sakaluk, S.K. 2008. The effect of age and previous mating experience on pre- and post-copulatory mate choice in female house crickets (Acheta domesticus L). Journal of Insect Behavior, 21, 203-212.

Adaptive Plasticity in Wild Field Crickets’ Acoustic Signaling

Adaptive Plasticity in Wild Field Crickets’ Acoustic Signaling

A new article in PLOS ONE written by

Susan Bertram, Sarah Harrison, Ian Thomson, and Lauren Fitzsimmons

Click here for a copy of our PLOS ONE article

Click here for the Bertram Lab website

http://science.nature.nps.gov/im/units/ucbn/monitor/sagegrouse/sagegrouse.cfm

Figure 1: Changes to the density of competitors is an example of environmental variability commonly faced by organisms in the wild.

Environmental change is a common obstacle that animals have to overcome. Temporal or spatial changes in the weather, food availability, predator density, mate availability, or competitor density are just some examples of the environmental variability commonly faced by animals in the wild (Figure 1).The optimal phenotype (an organism’s morphology, physiology, behavior etc.) for a given environment often changes as environmental conditions change. For example, a caterpillar feeding on oak flowers will develop into a mimic of an oak catkin, while a sibling caterpillar feeding on leaves will develop into a mimic of a twig (Figure 2). These caterpillars may avoid predation by camouflaging themselves to match their environments.

Photo source: Whitman and Agrawal

Figure 2: Nemoria arizonaria caterpillars: (a) summer broods feed on oak leaves and
Develop to resemble oak twigs, (b) while spring broods feed on and resemble oak catkins (Whitman and Agrawal 2009).

Figure 3: Hydrangea macrophylla flowers grown in acidic soil produces blue flowers, while flowers grown in neutral or alkaline soil produce pink flowers.

Figure 3: Hydrangea macrophylla flowers grown in acidic soil produces blue flowers, while flowers grown in neutral or alkaline soil produce pink flowers.

If individuals are unable to adapt to changes in their surroundings, they may experience reduced survival or reproductive success compared to individuals who are able to adapt.Phenotypic plasticity is the ability to change behaviour, appearance, or other characteristics in response to environmental change. For instance, Hydrangea macrophylla produce either blue or pink flowers depending on soil alkalinity (Figure 3). Phenotypic plasticity may be adaptive as long as environmental variation is predictable and the benefits of plasticity outweigh the costs. For instance, phenotypic plasticity in sexually selected traits may be particularly adaptive if there are frequent predictable changes to the socio-sexual environment (e.g. fluctuation in mate density).

In our study, recently published in PLOS ONE, we examined the phenotypic plasticity of male field crickets’ sexually selected acoustic signals for two species: the fall field cricket, Gryllus pennsylvanicus and the spring field cricket, G. veletis (Figure 4).

Figure 3: The fall field cricket (Left- G. pennsylvanicus male) and the spring field cricket (Right- G. veletis male) are both found throughout North America.

Figure 4: The fall field cricket (Left- G. pennsylvanicus male) and the spring field cricket (Right- G. veletis male) are both found throughout North America.

Male crickets produce acoustic signals by raising their forewings and rubbing them together, each closing stroke producing a pulse of sound with 3-4 pulses grouped into a chirp (Figure 5). These acoustic calls are used to attract females, and females choose mates on the basis of the quantity and fine-scale structure of these signals. Female movement and sexual receptivity in the wild often follows predictable fluctuations throughout the day (diel rhythms), which may partially be explained by hourly fluctuations in the biotic (predator and parasite density) and abiotic (light and temperature levels) environment. Therefore, males that synchronize their daily signalling rhythms to match female mating activity are likely to have higher reproductive success. Here we examined how male acoustic signalling in the laboratory changes over the course of the day and compared this to the known reports of diel rhythms in female mating activity in the wild.
We captured wild males and brought them to our laboratory at Carleton University where we recorded their acoustic signalling for 2-4 days. We found that both spring and fall field crickets exhibited phenotypically plastic signalling behavior, with most males signalling more often and more attractively during the time of day when female mating activity is the highest in the wild (fall field cricket: night time and early morning; spring field cricket: early morning and afternoon). Most male crickets (from both species) chirped more often during the time of the day that female mating activity is highest in the wild (Figure 5). However, a few males of each species signalled in a seemingly maladaptive manner during times when female mating activity is lowest in the wild. Diel rhythms in signalling also differed across species, which may be beneficial in preventing lethal hybridization when spring and fall field crickets overlap temporally and spatially in mid-summer.

Figure 4: Waveforms of long-distance mate attraction signals of one G. pennsylvanicus and one G. veletis male.  Figures show typical long-distance mate attraction signal for each species and how signaling typically changes during time periods indicative of low (A & B) and high (C & D) mating activity in the wild. Signal fine-scale properties are indicated as follows: ChD = chirp duration; IChD= interchirp duration; PPCh = pulses per chirp; PD = pulse duration; IPD = interpulse duration; and PP = pulse period, which combines PD and IPD.

Figure 4: Waveforms of long-distance mate attraction signals of one G. pennsylvanicus and one G. veletis male. Figures show typical long-distance mate attraction signal for each species and how signaling typically changes during time periods indicative of low (A & B) and high (C & D) mating activity in the wild. Signal fine-scale properties are indicated as follows: ChD = chirp duration; IChD= interchirp duration; PPCh = pulses per chirp; PD = pulse duration; IPD = interpulse duration; and PP = pulse period, which combines PD and IPD.

Overall, our findings suggest that male field crickets exhibit phenotypically plastic mate attraction signals and that diel rhythms in these signals are synchronized so that they are in phase with diel rhythms in female mating activity, suggesting that signalling plasticity may be adaptive. We have yet to determine how the costs and benefits of the phenotypic plasticity we observed in our species of field cricket ultimate affect fitness.

References:

Whitman, D. W. and A. A. Agrawal. What is Phenotypic Plasticity and why is it Important? Pages 1-63 in: D. W. Whitman and T. N. Ananthakrishna (editors), Phenotypic plasticity of insects: Mechanisms and consequences. Science Publishers, Inc, Enfield, NH

Male crickets change their behaviour when watched by an audience

Playing to an audience: The social environment influences aggression and victory displays

A new article in Biology Letters by Lauren Fitzsimmons and Susan Bertram

Click here for a copy of our Biology Letters article

Click here for the Bertram Lab website

Click here for Lauren’s website

http://wallpaperscraft.com/download/stag_beetle_fight_male_female_36738

Figure 1. Two male stag beetles fight while a female watches

In nature, many animals fight for dominance, territories, and mates, and fights often occur when others are watching (Figure 1). For example, eavesdropping fish are more likely to initiate fights with a loser than a winner, imposing an immediate cost to the loser. Fighters also sometimes change their behaviour when they know they are being watched.

Figure 2. Football players are famous for their victory celebrations after a touchdown.

Figure 2. Football players are famous for their victory celebrations after a touchdown.

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Humans do this regularly: consider the classic victory dance following a touchdown in a football game (Figure 2). Fish do this too: male fish have been shown to change their female preference in the presence of a potential competitor.

We conducted the first investigation of how audiences affect fighting and victory displays in an invertebrate, the spring field cricket (Gryllus veletis; Figure 3). Male field crickets frequently engage in fights over resources (see video here).

Winning a fight increases a male’s mating success through dominance which provides access to mate attraction territories, and because females usually prefer to mate with dominant males. Cricket fight winners often advertise their success using victory displays of aggressive songs and body jerks (shown after the fight in the video linked above). Because cricket densities can be high in field crickets, and mate attraction and fights occur in close proximity, many fights are likely to occur with female and male audiences nearby.

Figure 3. Two male spring field crickets (Gryllus veletis) engaged in a fight (photo credit: Louis Gagnon).

Figure 3. Two male spring field crickets (Gryllus veletis) engaged in a fight (photo credit: Louis Gagnon).

We investigated aggression and victory display behaviour in both field-captured and laboratory-reared crickets to explore the effect of rearing environment on these behaviours. We predicted that males with social experience would change their behaviour depending on the social context, whereas inexperienced lab-reared males would be less likely to respond to the presence of an audience. We found that the type of audience and the rearing environment (field or lab) were important predictors of how males behaved during and after fights. Field-captured winners were more aggressive than laboratory-reared winners in the presence of an audience (Figure 4).

Figure 4. Field-captured males are more aggressive with an audience present than with no audience, while lab-reared males are less responsive to the social environment. Field-captured males are also more aggressive than lab-reared males when an audience is present but similar when no audience is present.

Figure 4. Field-captured males are more aggressive with an audience present than with no audience, while lab-reared males are less responsive to the social environment. Field-captured males are also more aggressive than lab-reared males when an audience is present but similar when no audience is present.

Field-captured winners produced more victory displays in the presence of a male audience compared to no audience, whereas the victory display behaviour of lab-reared males was similar across audiences and highly variable among males within audience conditions (Figure 5). Our results suggest that field-captured winners, in particular, dynamically adjust their fighting behaviour to potentially gain a reproductive benefit via female eavesdropping and may deter future aggression from rivals by advertising their aggressiveness and victories.

Female mating decisions and male fighting decisions may be influenced by information communicated during contests, and females may represent a valuable resource for the winner, providing advantages to elevated aggression by winners during fights. Indeed, we found that contest winners elevate aggression in the presence of a female audience compared to no audience. Contest winners may be selected to be more aggressive and give more victory displays with male audiences because these displays reduce the likelihood of future contests; victory displays may also advertise additional energy that could be used against potential rivals. It is unknown whether cricket audiences gain information through eavesdropping, but our results suggest potential payoffs for both victorious males and eavesdroppers.

Figure 5. Field-captured males elevate victory display behaviour when a male audience is present, indicating a potential browbeating function (boasting their victory to other males to deter future aggression).

Figure 5. Field-captured males elevate victory display behaviour when a male audience is present, indicating a potential browbeating function (boasting their victory to other males to deter future aggression).

The relative lack of response to audience treatments for lab-reared males may reflect a lack of social experience. Our findings suggest that experiments on naive lab-reared individuals may not accurately reflect the behaviour of wild animals in nature, and add to evidence that social experience is important in shaping the development of dynamic behaviours. Field-captured males may be better able to adjust their dynamic behaviours to different social environments after experiencing both their own aggressive encounters and observing interactions between other males.

Our study provides the first evidence that invertebrates modify their contest behaviour in the presence of an audience. The ability to perceive the presence and sex of an audience and adjust behaviour accordingly is thus not restricted to vertebrates and may be more common across animals than previously recognized. Our study also provides the first evidence of an audience effect on victory display behaviour, and highlights the importance of the social environment in shaping animal behaviour.

Dr. Lauren Fitzsimmons

Dr. Lauren Fitzsimmons

ResearchBlogging.orgFitzsimmons, L.P. and S.M. Bertram (2013). Playing to an audience: The social environment influences aggression and victory displays. Biology Letters

Deceptive Males: Not All Sexual Signals Are Honest

Calling, Courtship, and Condition in the Fall Field Cricket, Gryllus pennsylvanicus

A new article in PLOS ONE written by

Sarah Harrison, Ian Thomson, Caitlin Grant, and Susan Bertram

Click here for a copy of our PLOS ONE article

Click here for a press release by Carleton University

Click here for the Bertram Lab website

Acoustic and physiological sexual signals

Figure 1. Acoustic and physiological sexual signals

Males often produce signals to attract females to mate. These sexual signals come in many forms: acoustic (e.g. calls of songbirds and crickets), visual (e.g. deer antlers, bright colourful bird plumage, colourful tropical fish scales; Figure 1), and chemical (e.g. pheromones). These signals are thought to provide females with information about male quality and potential mating benefits to help them choose between different males.

IMG_20111223_113319

Figure 2. Bright plumage and plump body size in a crested partridge.

For instance, if bright colourful bird plumage is strongly dependent on nutrition, then feather colour might signal that a male is healthy and good at finding high quality food (Figure 2). Mating with males that have colourful plumage might benefit females directly if males share their food, or indirectly if they pass on good genes for health and foraging ability to their offspring. Since poor quality males should be incapable of cheating and producing sexual signals that rival those of high quality males, sexual signals are thought to honestly indicate mating benefits. But are all sexual signals honest, or do males sometimes cheat when trying to attract a mate?

midlife-crisis-1

Figure 3. Are you compensating for something?
Source: http://www.blogger.com/img/blank.gif

Males might be more likely to produce dishonest signals (cheat) if they have a bleak reproductive future. For instance, starving, diseased, old, or dying males might invest all of their remaining energy into attracting females at the risk of dying sooner (Figure 3). This alternative idea, known as the terminal investment hypothesis, suggests that poor quality males might sometimes be able to produce signals that rival the signals of high quality males. These poor quality males would not, however, be able to provide females with the mating benefits that they were dishonestly advertizing.

We questioned whether all sexual signals are honest using fall field crickets (Gryllus pennsylvanicus). These crickets are commonly found throughout much of the U.S. and southern Canada. Males produce two types of mating calls by rubbing their hind wings together: one they use to attract females from a distance, the other they use to court attracted females. Both mate attraction and courtship calls are made up of a series of sound pulses that they group into chirps (Figure 4).

Cricket chirps

Figure 4. Sonograms (top) and waveforms (bottom) of a G. pennsylvanicus long-distance mate attraction call (A & B) and a courtship call (C & D).

We break down these chirps into smaller pieces (call rate, chirp rate, pulse rate, tick rate, number of pulses per chirp etc) to study them. Males can invest more or less energy into calling by changing the temporal properties of these pieces. Higher condition males should have more energy to devote to signalling than low condition males (‘Condition’ is an individual’s ability to find food and turn it into useable energy). Females should benefit from using calls to find mates because high condition males should pass on good genes for condition to their offspring.

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Figure 5. Size variation in G. pennsylvanicus

We recorded each male’s long-distance calls and courtship calls (click on the calls for examples), broke these calls down to their smaller temporal components, and then examined relationships between these temporal components and two different estimates of male condition: 1) body size (i.e. height; Figure 5) and 2) plumpness (i.e. fat).

We found evidence that males both honestly and dishonestly signal their condition. Male mate attraction calls honestly indicated body size, except that small males called with faster chirp rates than large males. Courtship calls dishonestly signalled male plumpness, as lean males produced calls with pulse and chirp rates similar to plump males, with males of intermediate plumpness having the lowest pulse and chirp rates. It costs more energy to call at faster pulse and chirp rates, so faster rates should be harder for low condition males. Our findings reveal that small and lean males cheat when calling to females: they call at equivalent or higher rates than large or plump males, suggesting not all sexual signals are honest.

Blog post “Deceptive Males: Not All Sexual Signals Are Honest” was written by Sarah Harrison, a PhD candidate in the Bertram Lab.

Sarah J. Harrison

Sarah J. Harrison

ResearchBlogging.orgHarrison SJ, Thomson IR, Grant CM, & Bertram SM (2013). Calling, Courtship, and Condition in the Fall Field Cricket, Gryllus pennsylvanicus. PloS one, 8 (3) PMID: 23527313