Using the <em>FishSim</em> Animation Toolchain to Investigate Fish Behavior: A Case Study on Mate-Choice Copying In Sailfin Mollies

Over the last decade, employing computer animations for animal behavior research has increased due to its ability to non-invasively manipulate the ...

University of Siegen

Virtual is the new reality.

In animals, including humans, the social environment can serve as a public information network in which individuals can gather public information about the quality of potential mates by observing conspecifics during sexual interactions. The observing individual itself is also a part of this information network. When recognized by the observed conspecifics as an audience, his/her presence could influence the sexual interaction between those individuals, because the observer might be considered as a potential mate or competitor. One of the most challenging questions in sexual selection to date is how the use of public information in the context of mate choice is linked to the fitness of individuals. Here, we could show that public information influences mate-choice behaviour in sailfin molly males, Poecilia latipinna, and influences the amount of sperm males transfer to a female partner. In the presence of an audience male, males spent less time with the previously preferred, larger of two females and significantly more time with the previously non-preferred, smaller female. When males could physically interact with a female and were faced with an audience male, three audience females or no audience, males transferred significantly more sperm to a female partner in the presence of an audience male than with female audience or no audience and spent less time courting his female partner. This is the first study showing that public information use turns into fitness investment, which is the crucial factor to understand the role of public information in the dynamic processes in sexual selection.

Public information influences sperm transfer to females in sailfin molly males.

Biodiversity is rapidly declining globally. One strategy to help to conserve species is to breed species in captivity and release them into suitable habitats. The way that reintroduced animals explore new habitats and/or disperse from the release site is rarely studied in detail and represents key information for the success of reintroduction projects. The European bison (Bison bonasus L. 1758) was the largest surviving herbivore of the post-glacial megafauna in Europe before it became extinct in the wild, surviving only in captivity since 1919. We investigated the exploration behavior of a herd of European bison reintroduced into the Rothaargebirge, a commercial forest in low range mountain intensively used and densely populated by humans, in the first six months after release. We focused on three questions: (1) how did the European bison move and utilize the habitat on a daily basis, (2) how did the animals explore the new environment, and (3) did their habitat preferences change over time. The European bison dispersed away from their previous enclosure at an average rate of 539 m/month, with their areas of daily use ranging from 70 to 173 ha, their movement ranging from 3.6 km to 5.2 km per day, and their day-to-day use of areas ranged between 389 and 900 m. We could identify three major exploration bouts, when the animals entered and explored areas previously unknown to them. During the birthing phase, the European bison reduced daily walking distances, and the adult bull segregated from the herd for 58 days. Around rut, roaming behavior of the herd increased slightly. The animals preferred spruce forest, wind thrown areas and grassland, all of which are food abundant habitat types, and they avoided beech forest. Habitat preference differed slightly between phases of the study period, probably due to phenological cycles. After six months, the complete summer home range was 42.5 km2. Our study shows that a small free-ranging herd of European bison can live in an area intensively used by humans and describes in detail the initial roaming behavior and habitat utilization of the animals. 

First Steps into the Wild - Exploration Behavior of European Bison after the First Reintroduction in Western Europe.

Animals observing conspecifics during mate choice can gain additional information about potential mates. However, the presence of an observer, if detected by the observed individuals, can influence the nature of the behavior of the observed individuals, called audience effect. In zebra finches (Taeniopygia guttata castanotis), domesticated males show an audience effect during mate choice. However, whether male and female descendants of the wild form show an audience effect during mate choice is unknown. Therefore, we conducted an experiment where male and female focal birds could choose between two distinctive phenotypes of the opposite sex, an artificially adorned stimulus bird with a red feather on the forehead and an unadorned stimulus bird, two times consecutively, once without an audience and once with an audience bird (same sex as test bird). Males showed an audience effect when an audience male was present and spent more time with adorned and less time with unadorned females compared to when there was no audience present. The change in time spent with the respective stimulus females was positively correlated with the time that the audience male spent in front of its cage close to the focal male. Females showed no change in mate choice when an audience female was present, but their motivation to associate with both stimulus males decreased. In a control for mate-choice consistency there was no audience in either test. Here, both focal females and focal males chose consistently without a change in choosing motivation. Our results showed that there is an audience effect on mate choice in zebra finches and that the response to a same-sex audience was sex-specific. 

Sex-Specific Audience Effect in the Context of Mate Choice in Zebra Finches.

Rapid technical advances in the field of computer animation (CA) and virtual reality (VR) have opened new avenues in animal behavior research. Animated stimuli are powerful tools as they offer standardization, repeatability, and complete control over the stimulus presented, thereby "reducing" and "replacing" the animals used, and "refining" the experimental design in line with the 3Rs. However, appropriate use of these technologies raises conceptual and technical questions. In this review, we offer guidelines for common technical and conceptual considerations related to the use of animated stimuli in animal behavior research. Following the steps required to create an animated stimulus, we discuss (I) the creation, (II) the presentation, and (III) the validation of CAs and VRs. Although our review is geared toward computer-graphically designed stimuli, considerations on presentation and validation also apply to video playbacks. CA and VR allow both new behavioral questions to be addressed and existing questions to be addressed in new ways, thus we expect a rich future for these methods in both ultimate and proximate studies of animal behavior.

Technical and conceptual considerations for using animated stimuli in studies of animal behavior.

Animal behavior researchers often face problems regarding standardization and reproducibility of their experiments. This has led to the partial substitution of live animals with artificial virtual stimuli. In addition to standardization and reproducibility, virtual stimuli open new options for researchers since they are easily changeable in morphology and appearance, and their behavior can be defined. In this article, a novel toolchain to conduct behavior experiments with fish is presented by a case study in sailfin mollies . As the toolchain holds many different and novel features, it offers new possibilities for studies in behavioral animal research and promotes the standardization of experiments. The presented method includes options to design, animate, and present virtual stimuli to live fish. The designing tool offers an easy and user-friendly way to define size, coloration, and morphology of stimuli and moreover it is able to configure virtual stimuli randomly without any user influence. Furthermore, the toolchain brings a novel method to animate stimuli in a semiautomatic way with the help of a game controller. These created swimming paths can be applied to different stimuli in real time. A presentation tool combines models and swimming paths regarding formerly defined playlists, and presents the stimuli onto 2 screens. Experiments with live sailfin mollies validated the usage of the created virtual 3D fish models in mate-choice experiments.

The virtual lover: variable and easily guided 3D fish animations as an innovative tool in mate-choice experiments with sailfin mollies-I. Design and implementation.

The use of computer animation in behavioral research is a state-of-the-art method for designing and presenting animated animals to live test animals. The major advantages of computer animations are: (1) the creation of animated animal stimuli with high variability of morphology and even behavior; (2) animated stimuli provide highly standardized, controlled and repeatable testing procedures; and (3) they allow a reduction in the number of live test animals regarding the 3Rs principle. But the use of animated animals should be attended by a thorough validation for each test species to verify that behavior measured with live animals toward virtual animals can also be expected with natural stimuli. Here we present results on the validation of a custom-made simulation for animated 3D sailfin mollies  and show that responses of live test females were as strong to an animated fish as to a video or a live male fish. Movement of an animated stimulus was important but female response was stronger toward a swimming 3D fish stimulus than to a "swimming" box. Moreover, male test fish were able to discriminate between animated male and female stimuli; hence, rendering the animated 3D fish a useful tool in mate-choice experiments with sailfin mollies.

The virtual lover: variable and easily guided 3D fish animations as an innovative tool in mate-choice experiments with sailfin mollies-II. Validation.

The louse fly  is an obligate blood-sucking ectoparasite of the common swift  As a result of reduction of the wings,  is unable to fly; thus, an effective and reliable attachment to their host's plumage is of utmost importance. The attachment system of  shows several modifications in comparison to that of other calyptrate flies, notably the large tridentate claws and the dichotomously shaped setae located on the pulvilli. Based on data from morphological analysis, confocal laser scanning microscopy, cryo-scanning electron microscopy and attachment force experiments performed on native (feathers) as well as artificial substrates (glass, epoxy resin and silicone rubber), we showed that the entire attachment system is highly adapted to the fly's lifestyle as an ectoparasite. The claws in particular are the main contributor to strong attachment to the host. Resulting attachment forces on feathers make it impossible to detach  without damage to the feathers or to the legs of the louse fly itself. Well-developed pulvilli are responsible for the attachment to smooth surfaces. Both dichotomously shaped setae and high setal density explain high attachment forces observed on smooth substrates. For the first time, we demonstrate a material gradient within the setae, with soft, resilin-dominated apical tips and stiff, more sclerotized bases in Diptera. The empodium seems not to be directly involved in the attachment process, but it might operate as a cleaning device and may be essential to maintain the functionality of the entire attachment system.

Holding tight to feathers - structural specializations and attachment properties of the avian ectoparasite  (Diptera, Hippoboscidae).

Animals often use public information for mate-choice decisions by observing conspecifics as they choose their mates and then copying this witnessed decision. When the copier, however, is detected by the choosing individual, the latter often alters its behavior and spends more time with the previously non-preferred mate. This behavioral change is called the audience effect. The deception hypothesis states that the choosing individual changes its behavior to distract the audience from the preferred mate. The deception hypothesis, however, only applies if the audience indeed copies the pretended mate choice of the observed individual. So far, this necessary prerequisite has never been tested. We investigated in Atlantic molly males and females whether, first, focal fish show an audience effect, i.e., alter their mate choices in the presence of an audience fish, and second, whether audience fish copy the mate choice of the focal fish they had just witnessed. We found evidence that male and female Atlantic mollies copy the pretended mate choice of same-sex focal fish. Therefore, a necessary requirement of the deception hypothesis is fulfilled. Our results show that public information use in the context of mate choice can be costly.

Klaudia Witte

Research Group of Ecology and Behavioral Biology,
Institute of Biology,
Research Group of Ecology and Behavioral Biology, Institute of Biology

University of Siegen

Using the FishSim Animation Toolchain to Investigate Fish Behavior: A Case Study on Mate-Choice Copying In Sailfin Mollies

Klaudia Witte

.css-o250i3{font-size:18px;font-family:Open Sans,sans-serif;line-height:21.6px;}Research Group of Ecology and Behavioral Biology,Institute of Biology,Research Group of Ecology and Behavioral Biology, Institute of Biology

University of Siegen

Using the FishSim Animation Toolchain to Investigate Fish Behavior: A Case Study on Mate-Choice Copying In Sailfin Mollies

Research Group of Ecology and Behavioral Biology,
Institute of Biology,
Research Group of Ecology and Behavioral Biology, Institute of Biology