New paper out in Biology Letters

Today my latest paper came out in Biology Letters! You can find it here.

The spectacular and complex visual patterns created by animal groups moving together have fascinated humans since the beginning of time. Think of the highly synchronized movements of a flock of starlings, or the circular motion of a school of barracudas. Using state-of-the-art robotics, a research team from the University of Konstanz, Science of Intelligence, and the Leibniz Institute of Freshwater Ecology and Inland Fisheries (IGB) shows that animals’ speed is fundamental for collective behavioral patterns, and that ultimately it is the faster individuals that have the strongest influence on group-level behavior. The study, published in Biology Letters of the Royal Society, gives new insights on complex collective behavioral patterns in nature, and provides knowledge that could help develop robotic systems that move collectively, such as robot swarms, driverless cars, and drones.

Researchers have long focused on identifying the emergence of collective patterns. Thanks to a combination of behavioral experiments, computer simulations, and field observations, it is clear that many seemingly complex patterns can actually be explained by relatively simple rules: move away from others if they get too near, speed up towards others if they get too far away, and otherwise move at the same speed and align with your group mates.

Besides understanding the rules that individuals follow when interacting with others, we need to consider the behaviors and characteristics of those individuals that make up the group and determine their influence for collective outcomes” says Dr. Jolle Jolles, a scientist at the Zukunftskolleg, University of Konstanz, and lead author of the study. “Across the animal kingdom, it has been found again and again that animals tend to differ considerably from one another in their behavior such as in terms of their activity, risk-taking, and social behavior“. What are the consequences of this behavioral heterogeneity when it comes to collective behavior? And how can one test for its social consequences?

To disentangle the role of individual differences in collective behavior and the mechanisms underlying this type of behavior, the research team built “Robofish”, a robotic fish that not only realistically looks and behaves like a guppy – a small tropical freshwater fish – , but also interacts with the live fish in a natural way. The experimenters paired the robotic fish with a guppy and programmed it to always follow its partner and copy its movements, lacking however any movement preferences of its own. The team then used high-definition video tracking and a closed-loop feedback system to let the robotic fish respond to the live fish’s actions in real-time.

One of Robofish’s simple interaction rules was to keep a constant distance to its shoal mate” explains Dr. David Bierbach, who works within the Berlin-based Excellence Cluster ‘Science of Intelligence’ at the HU Berlin and the IGB, and is senior author on the paper. “Using this rule, our Robofish tried to keep the same distance to the live fish by accelerating and decelerating whenever the live fish did. Also, programming the robotic fish without any own movement preferences gave us the unique opportunity to investigate how individual differences in the behavior of the live fish led to group-level differences. In short, with our unique approach, we could isolate the effect of the fish’s movement speed on the pair’s collective behavior“.

The researchers first quantified the guppies’ natural movement speed by observing their movements when alone in an open environment, and found that there were large individual differences in how fast guppies tended to move. When the fish were subsequently tested with Robofish, the fish and Robofish tended to swim naturally together as a pair. However, the researchers observed that there were large differences in the social behaviors between the pairs: pairs in which the guppy had a faster movement speed tended to be much more aligned, more coordinated, and less cohesive, and the guppy emerged as a clearer leader. As Robofish behaved according to the same identical rules with each and every guppy, it is the individual speed of the guppies that must have led to these differences in group-level properties.

By involving state-of-the-art robotics, this research shows that individual speed is a fundamental factor in the emergence of collective behavioral patterns. As individual differences in speed are associated with a broad range of phenotypic traits among grouping animals, such as their size, age, and hunger level, the results of this study may help understand the role of such heterogeneity in animal groups.

Future studies using the interactive Robofish will focus on other aspects of collective behavior: For example, how can animals act in synchrony if they just respond to the actions of their neighbors? “We want to improve Robofish’s software so that it can predict and anticipate the live fish’s next steps, which is assumed to be how animals do it.” says David Bierbach.

Understanding these mechanisms is not only fundamentally important as it reveals information about the mechanisms that underlie collective behavior and decisions, but also because this knowledge can be applied to artificial systems and used to develop machines that move collectively, such as robot swarms, driverless cars, and drones.

New preprint about robofish and group-behaviour

Today I released a new preprint on bioRxiv, Group-level patterns emerge from individual speed as revealed by an extremely social robotic fish, which is the result of a great collaboration with David Bierbach and colleagues at the Humboldt Universität zu Berlin.

In this paper we present results of an experiment to investigate how the speed of individual group members leads to group-level patterns. We paired guppies with a biomimetic robot that was programmed to always follow and lack any individual preferences of its own. We used a state-of-the art closed-loop tracking and feedback system to be able to properly control for the influence of individual heterogeneity of the individual’s group members.

We show that individual differences in guppies’ movement speed were highly repeatable and shaped key collective patterns: higher individual speeds resulted in stronger leadership, lower cohesion, higher alignment, and better temporal coordination in the pairs. By combining the strengths of individual-based models and observational work with state-of-the-art robotics, we provide novel evidence that individual speed is a key, fundamental process in the emergence of collective behaviour.

On the front cover of TREE!

I am excited to say that our review in Trends in Ecology and Evolution, after already being available online, is out now in print and is shining on the front cover! I took this photo of this stunning stickleback school while snorkelling in the Bodensee to study their collective behaviour. Read our open access paper here.

New paper out on personality, plasticity, and predictability

Today my latest paper came out in Animal Behaviour, one of my favourite journals. It is titled “Personality, plasticity and predictability in sticklebacks: bold fish are less plastic and more predictable than shy fish“. In this paper, which is a result of a collaboration with Neeltje Boogert and Yimen Araj-Ayoy and a MSc project of Helen Briggs, we present an extensive experimental study focused on better understanding the sources of behavioural variation among individual animals.

In short, we tested 80 three-spined sticklebacks repeatedly on their boldness across a 10-week testing period and automatically tracked their movements. We then employed advanced statistical model techniques (GLMMs and DHGLMs) to use this large behavioural dataset to investigate the potential links between the personality (consistent differences in average behaviour), the plasticity (how individuals change their behaviour over time/contexts), and predictability (the remaining intra-individual variation after accounting for personality and plasticity differences) in behaviour.

Besides detecting large consistent individual differences in boldness and the extent to which fish changed this behaviour over time (temporal plasticity), we found that boldness personality and plasticity were negatively linked, with bold fish changing little in their behaviour over time. Interestingly, there were still large individual differences in the remaining behavioural variation, with bold fish showing much less behavioural variation and thus behaving more predictable than shy fish. Importantly, these results suggest that boldness, plasticity and predictability may be fundamentally linked and form part of the same behavioural syndrome.

Jolles, J. W., Briggs, H. D., Araya-Ajoy, Y. G., & Boogert, N. J. (2019). Personality, plasticity and predictability in sticklebacks: bold fish are less plastic and more predictable than shy fish. Animal Behaviour, 154, 193–202.

The paper, which was one of the Editor’s Choice papers for August 2019, can be found here.

New paper out in Proceedings of the Royal Society B!

From swarm to school, stickleback groups differ repeatedly in their collective performance  

 among schooling fish, groups can have different collective personalities, with some shoals sticking closer together, being better coordinated, and showing clearer leadership than others.

For centuries, scientists and non-scientists alike have been fascinated by the beautiful and often complex collective behaviour of animal groups, such as the highly synchronised movements of flocks of birds and schools of fish. Often, those spectacular collective patterns emerge from individual group members using simple rules in their interactions, without requiring global knowledge of their group.

In recent years it has also become apparent that, across the animal kingdom, individual animals often differ considerably and consistently in their behaviour, with some individuals being bolder, more active, or more social than others.

New research conducted at the University of Cambridge’s Department of Zoology suggests that observations of different groups of schooling fish could provide important insights into how the make-up of groups can drive collective behaviour and performance.

In the study, published today in the journal Proceedings of the Royal Society B, the researchers created random groups of wild-caught stickleback fish and subjected them repeatedly to a range of environments that included open spaces, plant cover, and patches of food.

Read further…

New paper out in Current Biology!

My latest paper on the collective behaviour of stickleback shoals is out today in the journal Current Biology!

Jolles, JW, Boogert, NJ, Sridhar, VH, Couzin, ID, Manica, A. (2017) Consistent individual differences drive collective behaviour and group functioning of schooling fish. Current Biology 27: 1-7. doi: 10.1016/j.cub.2017.08.004 (link).

Highly coordinated school of three-spined sticklebacks swimming in the blue waters of the Bodensee near Konstanz, Southern Germany. Photo: Jolle W. Jolles

Highly coordinated school of three-spined sticklebacks swimming in the blue waters of the Bodensee near Konstanz, Southern Germany. Photo: Jolle W. Jolles

New research sheds light on how “animal personalities” – inter-individual differences in animal behaviour – can drive the collective behaviour and functioning of animal groups such as schools of fish, including their cohesion, leadership, movement dynamics, and group performance. These research findings from the University of Konstanz, the Max Planck Institute of Ornithology and the University of Cambridge provide important new insights that could help explain and predict the emergence of complex collective behavioural patterns across social and ecological scales, with implications for conservation and fisheries and potentially creating bio-inspired robot swarms. It may even help us understand human society and team performance. The study is published in the 7 September 2017 issue of Current Biology.

Read further…

Bold fish are less sociable and thereby affect leadership and group coordination

Recent research of colleagues and I at the University of Cambridge has revealed that sticklebacks with bolder personalities are not only better leaders but also less sociable than more timid fish. The behaviour of these bolder fish shapes the dynamics of the group.

See a 4min video about our research here a 4min video about the paper here:

Throughout the animal kingdom, individuals often live and move together in groups, from swarms of insects to troops of primates. Individual animals may benefit from being part of groups, which provide protection from predators and help in finding food. To ensure that individuals reap the benefits of togetherness, group members coordinate their behaviour. As a result, leaders and followers emerge.

Within groups, animals differ from each other in how they cope with their environment and often exhibit distinctive traits, such as boldness or sociability. Even three-spined sticklebacks, the ‘tiddlers’ collected from streams and ponds by generations of schoolchildren, can be described in terms of their personalities: some are bolder and take more risks, while others are more timid and spend more of their time hiding in the weeds.

Research carried out in the Zoology Department at the University of Cambridge suggests that observations of these tiny fish, and how they interact with one another, could provide important insights into the dynamics of social groups, including humans.

Jolle Jolles, lead author of the study, said: “Although we now know that the spectacular collective behaviours we find throughout the animal kingdom can often be explained by individuals following simple rules, little is known about how this may be affected by the personality types that exist within the group.

Experimental design. Fish were tested twice for one hour in the risk-taking task during two subsequent sessions. During the pairing session fish could see and interact with one another through a transparent partition.

Experimental design. Fish were tested twice for one hour in the risk-taking task during two subsequent sessions. During the pairing session fish could see and interact with one another through a transparent partition.

“Our research shows that personality plays an important role in collective behaviour and that boldness and sociability may have significant, and complementary, effects on the functioning of the group.”

In the study, the researchers studied the behaviours of sticklebacks in tanks containing gravel and weed to imitate patches of a riverbed. The tanks were divided into two lanes by transparent partitions and randomly-selected pairs of fish were placed one in each lane. Separated by the see-through division, the fish were able to see and interact with one another.


The positions and movements of the individual sticklebacks were recorded using sophisticated tracking technology, enabling accurate comparisons to be made of each fish’s role in the collective movement of the pair.

“We found that individuals differed considerably and consistently in their tendency to approach their partner,” said Jolles. The study showed that more sociable individuals tended to be coordinated in their behaviour while less sociable individuals were more inclined to lead.

Dr Andrea Manica, reader at the Department of Zoology and co-author of the paper, added: “Our research revealed that the tendency of fish to approach their partner was strongly linked to their boldness: bolder fish were less sociable than their more timid group mates.”

Jolles explains that sociability may form part of a broader behavioural syndrome. “Our results suggest that bolder, less sociable individuals may often lead simply because they are less reluctant to move away from their partners, whereas shyer, more sociable, individuals become followers because they prioritise staying close to others,” he said.


“Differences in boldness and sociability may be expressions of underlying risk-prone or risk-averse behavioural types, as risk-averse individuals may be more motivated to group together and to respond to other individuals in order to avoid predation.”

The findings of this study suggest that leadership and group coordination can be strongly affected by personality differences in the group and that boldness and sociability may play important but complementary roles in collective behaviour.

Jolles added: “Now we know these personality traits affect the collective movements of pairs of fish, the next step is to understand their role in the functioning and success of larger, more dynamic groups.”

See a 4min video in which we explain our paper in more detail below:

jolles-animal-behaviour-stickleback-leadershipClick here to download the paper.

Jolles JW, et al. (2015) The role of social attraction and its link with boldness in the collective movements of three-spined sticklebacks. Animal Behaviour, published online 2 Dec. Doi: 10.1016/j.anbehav.2014.11.004

Leadership in fish affected by previous experiences and linked to personality

Leadership behaviour is affected by social experiences from previous partners and depends on an individual’s personality, as shown by our latest study with three-spined stickleback fish, now published in Behavioral Ecology.


From the political affairs we see on the news, to making decisions with your friends, leadership is all around us. But next to humans, leaders and followers can also be found in many group-living animals, such as fish, birds, and primates.

Social animals may receive benefits from grouping such as protection from predators and help in finding food. But to ensure individuals reap the benefits of grouping, they must time and coordinate their behaviour with the emergence of potential leaders and followers as a result.
Read further…