Colin R. Twomey
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Research Subjects
Colin R. Twomey has a total of 27 co-authors. Here are the top co-authors by number of shared publications:
- Iain Douglas Couzin
Department of Collective Behaviour, Max Planck Institute of Animal Behavior, 78464 Konstanz, Germany
4 shared publications - David H. Brainard
Department of Psychology, University of Pennsylvania, Philadelphia, PA, USA
3 shared publications - Jacob D. Davidson
Department of Collective Behaviour, Max Planck Institute of Animal Behavior, 78464 Konstanz, Germany
3 shared publications - Joshua B. Plotkin
Department of Biology, University of Pennsylvania, Philadelphia, USA
3 shared publications Matthew M. G. Sosna
Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA
3 shared publications- Vivek Hari Sridhar
Department for the Ecology of Animal Societies, Max Planck Institute of Animal Behavior, Konstanz (78467), Germany
3 shared publications
Latest Publications
- Open Access
- 28 February 2024
- Published by National Academy of Sciences in Proceedings of the National Academy of Sciences
- Vol. 121 (10)
History constrains the evolution of efficient color naming, enabling historical inference
Color naming in natural languages is not arbitrary: It reflects efficient partitions of perceptual color space [T. Regier, P. Kay, N. Khetarpal,Proc. Natl. Acad. Sci. U.S.A.104, 1436–1441 (2007)] modulated by the relative needs to communicate about different colors [C. Twomey, G. Roberts, D. Brainard, J. Plotkin,Proc. Natl. Acad. Sci. U.S.A.118, e2109237118 (2021)]. These psychophysical and communicative constraints help explain why languages around the world have remarkably similar, but not identical, mappings of colors to color terms. Languages converge on a small set of efficient representations.But languages also evolve, and the number of terms in a color vocabulary may change over time. Here we show that history, i.e. the existence of an antecedent color vocabulary, acts as a nonadaptive constraint that biases the choice of efficient solution as a language transitions from a vocabulary of sizetoterms. Moreover, as efficient vocabularies evolve to include more terms they explore a smaller fraction of all possible efficient vocabularies compared to equally sized vocabularies constructed de novo. This path dependence of the cultural evolution of color naming presents an opportunity. Historical constraints can be used to reconstruct ancestral color vocabularies, allowing us to answer long-standing questions about the evolutionary sequences of color words, and enabling us to draw inferences from phylogenetic patterns of language change.- 28 February 2024
- Published by National Academy of Sciences in Proceedings of the National Academy of Sciences
- Vol. 121 (10)
Open AccessHistory constrains the evolution of efficient color naming, enabling historical inference
Color naming in natural languages is not arbitrary: It reflects efficient partitions of perceptual color space [T. Regier, P. Kay, N. Khetarpal,Proc. Natl. Acad. Sci. U.S.A.104, 1436–1441 (2007)] modulated by the relative needs to communicate about different colors [C. Twomey, G. Roberts, D. Brainard, J. Plotkin,Proc. Natl. Acad. Sci. U.S.A.118, e2109237118 (2021)]. These psychophysical and communicative constraints help explain why languages around the world have remarkably similar, but not identical, mappings of colors to color terms. Languages converge on a small set of efficient representations.But languages also evolve, and the number of terms in a color vocabulary may change over time. Here we show that history, i.e. the existence of an antecedent color vocabulary, acts as a nonadaptive constraint that biases the choice of efficient solution as a language transitions from a vocabulary of sizetoterms. Moreover, as efficient vocabularies evolve to include more terms they explore a smaller fraction of all possible efficient vocabularies compared to equally sized vocabularies constructed de novo. This path dependence of the cultural evolution of color naming presents an opportunity. Historical constraints can be used to reconstruct ancestral color vocabularies, allowing us to answer long-standing questions about the evolutionary sequences of color words, and enabling us to draw inferences from phylogenetic patterns of language change. - Open Access
Historical constraints on the evolution of efficient color naming
Color naming in natural languages is not arbitrary: it reflects efficient partitions of perceptual color space modulated by the relative needs to communicate about different colors. These psychophysical and communicative constraints help explain why languages around the world have remarkably similar, but not identical, mappings of colors to color terms. Languages converge on a small set of efficient representations. But languages also evolve, and the number of terms in a color vocabulary may change over time. Here we show that history, i.e. the existence of an antecedent color vocabulary, acts as a non-adaptive constraint that biases the choice of efficient solution as a language transitions from a vocabulary of size n to n+1 terms. Moreover, as vocabularies evolve to include more terms they explore a smaller fraction of all possible efficient vocabularies compared to equally-sized vocabularies constructed de novo. This path dependence on the cultural evolution of color naming presents an opportunity. Historical constraints can be used to reconstruct ancestral color vocabularies, allowing us to answer long-standing questions about the evolutionary sequences of color words, and enabling us to draw inferences from phylogenetic patterns of language change.
Open AccessHistorical constraints on the evolution of efficient color naming
Color naming in natural languages is not arbitrary: it reflects efficient partitions of perceptual color space modulated by the relative needs to communicate about different colors. These psychophysical and communicative constraints help explain why languages around the world have remarkably similar, but not identical, mappings of colors to color terms. Languages converge on a small set of efficient representations. But languages also evolve, and the number of terms in a color vocabulary may change over time. Here we show that history, i.e. the existence of an antecedent color vocabulary, acts as a non-adaptive constraint that biases the choice of efficient solution as a language transitions from a vocabulary of size n to n+1 terms. Moreover, as vocabularies evolve to include more terms they explore a smaller fraction of all possible efficient vocabularies compared to equally-sized vocabularies constructed de novo. This path dependence on the cultural evolution of color naming presents an opportunity. Historical constraints can be used to reconstruct ancestral color vocabularies, allowing us to answer long-standing questions about the evolutionary sequences of color words, and enabling us to draw inferences from phylogenetic patterns of language change.
- Open Access
- 20 February 2023
- Published by The Royal Society in Philosophical Transactions of the Royal Society B: Biological Sciences
- Vol. 378 (1874) , 20220062
Inferring social influence in animal groups across multiple timescales
Many animal behaviours exhibit complex temporal dynamics, suggesting there are multiple timescales at which they should be studied. However, researchers often focus on behaviours that occur over relatively restricted temporal scales, typically ones that are more accessible to human observation. The situation becomes even more complex when considering multiple animals interacting, where behavioural coupling can introduce new timescales of importance. Here, we present a technique to study the time-varying nature of social influence in mobile animal groups across multiple temporal scales. As case studies, we analyse golden shiner fish and homing pigeons, which move in different media. By analysing pairwise interactions among individuals, we show that predictive power of the factors affecting social influence depends on the timescale of analysis. Over short timescales the relative position of a neighbour best predicts its influence and the distribution of influence across group members is relatively linear, with a small slope. At longer timescales, however, both relative position and kinematics are found to predict influence, and nonlinearity in the influence distribution increases, with a small number of individuals being disproportionately influential. Our results demonstrate that different interpretations of social influence arise from analysing behaviour at different timescales, highlighting the importance of considering its multiscale nature. This article is part of a discussion meeting issue ‘Collective behaviour through time’.
- 20 February 2023
- Published by The Royal Society in Philosophical Transactions of the Royal Society B: Biological Sciences
- Vol. 378 (1874) , 20220062
Open AccessInferring social influence in animal groups across multiple timescales
Many animal behaviours exhibit complex temporal dynamics, suggesting there are multiple timescales at which they should be studied. However, researchers often focus on behaviours that occur over relatively restricted temporal scales, typically ones that are more accessible to human observation. The situation becomes even more complex when considering multiple animals interacting, where behavioural coupling can introduce new timescales of importance. Here, we present a technique to study the time-varying nature of social influence in mobile animal groups across multiple temporal scales. As case studies, we analyse golden shiner fish and homing pigeons, which move in different media. By analysing pairwise interactions among individuals, we show that predictive power of the factors affecting social influence depends on the timescale of analysis. Over short timescales the relative position of a neighbour best predicts its influence and the distribution of influence across group members is relatively linear, with a small slope. At longer timescales, however, both relative position and kinematics are found to predict influence, and nonlinearity in the influence distribution increases, with a small number of individuals being disproportionately influential. Our results demonstrate that different interpretations of social influence arise from analysing behaviour at different timescales, highlighting the importance of considering its multiscale nature. This article is part of a discussion meeting issue ‘Collective behaviour through time’.
- Top Cited PapersOpen Access
- 24 June 2022
- Published by American Association for the Advancement of Science (AAAS) in Science Advances
- Vol. 8 (25) , eabm6385
Subcritical escape waves in schooling fish
Theoretical physics predicts optimal information processing in living systems near transitions (or pseudo-critical points) in their collective dynamics. However, focusing on potential benefits of proximity to a critical point, such as maximal sensitivity to perturbations and fast dissemination of information, commonly disregards possible costs of criticality in the noisy, dynamic environmental contexts of biological systems. Here, we find that startle cascades in fish schools are subcritical (not maximally responsive to environmental cues) and that distance to criticality decreases when perceived risk increases. Considering individuals’ costs related to two detection error types, associated to both true and false alarms, we argue that being subcritical, and modulating distance to criticality, can be understood as managing a trade-off between sensitivity and robustness according to the riskiness and noisiness of the environment. Our work emphasizes the need for an individual-based and context-dependent perspective on criticality and collective information processing and motivates future questions about the evolutionary forces that brought about a particular trade-off.
- 24 June 2022
- Published by American Association for the Advancement of Science (AAAS) in Science Advances
- Vol. 8 (25) , eabm6385
Top Cited PapersOpen AccessSubcritical escape waves in schooling fish
Theoretical physics predicts optimal information processing in living systems near transitions (or pseudo-critical points) in their collective dynamics. However, focusing on potential benefits of proximity to a critical point, such as maximal sensitivity to perturbations and fast dissemination of information, commonly disregards possible costs of criticality in the noisy, dynamic environmental contexts of biological systems. Here, we find that startle cascades in fish schools are subcritical (not maximally responsive to environmental cues) and that distance to criticality decreases when perceived risk increases. Considering individuals’ costs related to two detection error types, associated to both true and false alarms, we argue that being subcritical, and modulating distance to criticality, can be understood as managing a trade-off between sensitivity and robustness according to the riskiness and noisiness of the environment. Our work emphasizes the need for an individual-based and context-dependent perspective on criticality and collective information processing and motivates future questions about the evolutionary forces that brought about a particular trade-off.
- Open Access
- 23 September 2021
- Published by National Academy of Sciences in Proceedings of the National Academy of Sciences
- Vol. 118 (39)
What we talk about when we talk about colors
Significance: Do we talk about some colors more often than others? And do the colors we communicate about most frequently vary across cultures? A classic finding shows that languages around the world partition colors into words in remarkably similar, although not identical, ways. The biology of human color perception helps explain similar color vocabularies across languages, but less is known about how often speakers need to reference different colors. The inference method we develop reveals extensive variation in communicative needs across colors, and a diversity in needs across 130 languages, which helps explain variation in their color vocabularies. Our results open the door to studying cross-cultural variation in demands on different colors, and factors that drive color demands in linguistic communities.
- 23 September 2021
- Published by National Academy of Sciences in Proceedings of the National Academy of Sciences
- Vol. 118 (39)
Open AccessWhat we talk about when we talk about colors
Significance: Do we talk about some colors more often than others? And do the colors we communicate about most frequently vary across cultures? A classic finding shows that languages around the world partition colors into words in remarkably similar, although not identical, ways. The biology of human color perception helps explain similar color vocabularies across languages, but less is known about how often speakers need to reference different colors. The inference method we develop reveals extensive variation in communicative needs across colors, and a diversity in needs across 130 languages, which helps explain variation in their color vocabularies. Our results open the door to studying cross-cultural variation in demands on different colors, and factors that drive color demands in linguistic communities.
- Open Access
- 7 July 2021
- Published by The Royal Society in Journal of The Royal Society Interface
- Vol. 18 (180) , 20210142
Collective detection based on visual information in animal groups
We investigate key principles underlying individual, and collective, visual detection of stimuli, and how this relates to the internal structure of groups. While the individual and collective detection principles are generally applicable, we employ a model experimental system of schooling golden shiner fish ( Notemigonus crysoleucas ) to relate theory directly to empirical data, using computational reconstruction of the visual fields of all individuals. This reveals how the external visual information available to each group member depends on the number of individuals in the group, the position within the group, and the location of the external visually detectable stimulus. We find that in small groups, individuals have detection capability in nearly all directions, while in large groups, occlusion by neighbours causes detection capability to vary with position within the group. To understand the principles that drive detection in groups, we formulate a simple, and generally applicable, model that captures how visual detection properties emerge due to geometric scaling of the space occupied by the group and occlusion caused by neighbours. We employ these insights to discuss principles that extend beyond our specific system, such as how collective detection depends on individual body shape, and the size and structure of the group.
- 7 July 2021
- Published by The Royal Society in Journal of The Royal Society Interface
- Vol. 18 (180) , 20210142
Open AccessCollective detection based on visual information in animal groups
We investigate key principles underlying individual, and collective, visual detection of stimuli, and how this relates to the internal structure of groups. While the individual and collective detection principles are generally applicable, we employ a model experimental system of schooling golden shiner fish ( Notemigonus crysoleucas ) to relate theory directly to empirical data, using computational reconstruction of the visual fields of all individuals. This reveals how the external visual information available to each group member depends on the number of individuals in the group, the position within the group, and the location of the external visually detectable stimulus. We find that in small groups, individuals have detection capability in nearly all directions, while in large groups, occlusion by neighbours causes detection capability to vary with position within the group. To understand the principles that drive detection in groups, we formulate a simple, and generally applicable, model that captures how visual detection properties emerge due to geometric scaling of the space occupied by the group and occlusion caused by neighbours. We employ these insights to discuss principles that extend beyond our specific system, such as how collective detection depends on individual body shape, and the size and structure of the group.
- Open Access
Collective detection based on visual information in animal groups
The spatio-temporal distribution of individuals within a group (i.e its internal structure) plays a defining role in how individuals interact with their environment, make decisions, and transmit information via social interactions. Group-living organisms across taxa, including many species of fish, birds, ungulates, and insects, use vision as the predominant modality to coordinate their collective behavior. Despite this importance, there have been few quantitative studies examining visual detection capabilities of individuals within groups. We investigate key principles underlying individual, and collective, visual detection of stimuli (which could include cryptic predators, potential food items, etc.) and how this relates to the internal structure of groups. While the individual and collective detection principles are generally applicable, we employ a model experimental system of schooling golden shiner fish ( Notemigonus crysoleucas ) to relate theory directly to empirical data, using computational reconstruction of the visual fields of all individuals to do so. Our integrative approach allows us to reveal how the external visual information available to each group member depends on the number of individuals in the group, the position within the group, and the location of the external visually-detectable stimulus. We find that in small groups, individuals have detection capability in nearly all directions, while in large groups, occlusion by neighbors causes detection capability to vary with position within the group. We then formulate a simple, and generally applicable, model that captures how visual detection properties emerge due to geometric scaling of the space occupied by the group and occlusion caused by neighbors. We employ these insights to discuss principles that extend beyond our specific system, such as how collective detection depends on individual body shape, and the size and structure of the group.
Open AccessCollective detection based on visual information in animal groups
The spatio-temporal distribution of individuals within a group (i.e its internal structure) plays a defining role in how individuals interact with their environment, make decisions, and transmit information via social interactions. Group-living organisms across taxa, including many species of fish, birds, ungulates, and insects, use vision as the predominant modality to coordinate their collective behavior. Despite this importance, there have been few quantitative studies examining visual detection capabilities of individuals within groups. We investigate key principles underlying individual, and collective, visual detection of stimuli (which could include cryptic predators, potential food items, etc.) and how this relates to the internal structure of groups. While the individual and collective detection principles are generally applicable, we employ a model experimental system of schooling golden shiner fish ( Notemigonus crysoleucas ) to relate theory directly to empirical data, using computational reconstruction of the visual fields of all individuals to do so. Our integrative approach allows us to reveal how the external visual information available to each group member depends on the number of individuals in the group, the position within the group, and the location of the external visually-detectable stimulus. We find that in small groups, individuals have detection capability in nearly all directions, while in large groups, occlusion by neighbors causes detection capability to vary with position within the group. We then formulate a simple, and generally applicable, model that captures how visual detection properties emerge due to geometric scaling of the space occupied by the group and occlusion caused by neighbors. We employ these insights to discuss principles that extend beyond our specific system, such as how collective detection depends on individual body shape, and the size and structure of the group.
- Open Access
What we talk about when we talk about color
Names for colors vary widely across languages, but color categories are remarkably consistent (Berlin & Kay, 1969). Shared mechanisms of color perception help explain consistent partitions of visible light into discrete color vocabularies (Regier et al., 2007). But the mappings from colors to words are not identical across languages, which may reflect communicative needs – how often speakers must refer to objects of different color (Gibson et al., 2017). Here we quantify the communicative needs of colors in 130 different languages by means of a novel inference algorithm. We find that communicative needs are not uniform: some regions of color space exhibit 30-fold greater demand for communication than other regions. The regions of greatest demand correlate with the colors of salient objects, including ripe fruits in primate diets. Our analysis also reveals a hidden diversity in the communicative needs of colors across different languages, which is partly explained by differences in geographic location and the local biogeography of linguistic communities. Accounting for language-specific, non-uniform communicative needs improves predictions for how a language maps colors to words, and how these mappings vary across languages. Our account closes an important gap in the compression theory of color naming, while opening new directions to study cross-cultural variation in the need to communicate different colors and its impact on the cultural evolution of color categories.
Open AccessWhat we talk about when we talk about color
Names for colors vary widely across languages, but color categories are remarkably consistent (Berlin & Kay, 1969). Shared mechanisms of color perception help explain consistent partitions of visible light into discrete color vocabularies (Regier et al., 2007). But the mappings from colors to words are not identical across languages, which may reflect communicative needs – how often speakers must refer to objects of different color (Gibson et al., 2017). Here we quantify the communicative needs of colors in 130 different languages by means of a novel inference algorithm. We find that communicative needs are not uniform: some regions of color space exhibit 30-fold greater demand for communication than other regions. The regions of greatest demand correlate with the colors of salient objects, including ripe fruits in primate diets. Our analysis also reveals a hidden diversity in the communicative needs of colors across different languages, which is partly explained by differences in geographic location and the local biogeography of linguistic communities. Accounting for language-specific, non-uniform communicative needs improves predictions for how a language maps colors to words, and how these mappings vary across languages. Our account closes an important gap in the compression theory of color naming, while opening new directions to study cross-cultural variation in the need to communicate different colors and its impact on the cultural evolution of color categories.
- Open Access
- 23 March 2020
- Published by Springer Nature in Theory in Biosciences
- Vol. 140 (4) , 361-377
Searching for structure in collective systems
From fish schools and bird flocks to biofilms and neural networks, collective systems in nature are made up of many mutually influencing individuals that interact locally to produce large-scale coordinated behavior. Although coordination is central to what it means to behave collectively, measures of large-scale coordination in these systems are ad hoc and system specific. The lack of a common quantitative scale makes broad cross-system comparisons difficult. Here we identify a system-independent measure of coordination based on an information-theoretic measure of multivariate dependence and show it can be used in practice to give a new view of even classic, well-studied collective systems. Moreover, we use this measure to derive a novel method for finding the most coordinated components within a system and demonstrate how this can be used in practice to reveal intrasystem organizational structure.
- 23 March 2020
- Published by Springer Nature in Theory in Biosciences
- Vol. 140 (4) , 361-377
Open AccessSearching for structure in collective systems
From fish schools and bird flocks to biofilms and neural networks, collective systems in nature are made up of many mutually influencing individuals that interact locally to produce large-scale coordinated behavior. Although coordination is central to what it means to behave collectively, measures of large-scale coordination in these systems are ad hoc and system specific. The lack of a common quantitative scale makes broad cross-system comparisons difficult. Here we identify a system-independent measure of coordination based on an information-theoretic measure of multivariate dependence and show it can be used in practice to give a new view of even classic, well-studied collective systems. Moreover, we use this measure to derive a novel method for finding the most coordinated components within a system and demonstrate how this can be used in practice to reveal intrasystem organizational structure.
- Open Access
A machine-vision approach for automated pain measurement at millisecond timescales
Objective and automatic measurement of pain in mice remains a barrier for discovery in both basic and translational neuroscience. Here we capture rapid paw kinematics during pain behavior in mice with high-speed videography and automated paw tracking with machine and deep learning approaches. Our statistical software platform, PAWS (Pain Assessment at Withdrawal Speeds), uses a univariate projection of paw position over time to automatically quantify fast paw dynamics at the onset of paw withdrawal and also lingering pain-related behaviors such as paw guarding and shaking. Applied to innocuous and noxious stimuli across six inbred mouse strains, a linear discriminant analysis reveals a two-dimensional subspace that separates painful from non-painful stimuli on one axis, and further distinguishes the severity of pain on the second axis. Automated paw tracking combined with PAWS reveals behaviorally-divergent mouse strains that display hypo- and hyper-sensitivity to mechanical stimuli. To demonstrate the efficacy of PAWS for detecting hypersensitivity to noxious stimuli, we chemogenetically activated pain-aversion neurons in the amygdala, which further separated the behavioral representation of pain-related behaviors along a low-dimensional path. Taken together, this automated pain quantification approach should increase the ease and objectivity of collecting rigorous behavioral data, and it is compatible with other neural circuit dissection tools for determining the mouse pain state.
Open AccessA machine-vision approach for automated pain measurement at millisecond timescales
Objective and automatic measurement of pain in mice remains a barrier for discovery in both basic and translational neuroscience. Here we capture rapid paw kinematics during pain behavior in mice with high-speed videography and automated paw tracking with machine and deep learning approaches. Our statistical software platform, PAWS (Pain Assessment at Withdrawal Speeds), uses a univariate projection of paw position over time to automatically quantify fast paw dynamics at the onset of paw withdrawal and also lingering pain-related behaviors such as paw guarding and shaking. Applied to innocuous and noxious stimuli across six inbred mouse strains, a linear discriminant analysis reveals a two-dimensional subspace that separates painful from non-painful stimuli on one axis, and further distinguishes the severity of pain on the second axis. Automated paw tracking combined with PAWS reveals behaviorally-divergent mouse strains that display hypo- and hyper-sensitivity to mechanical stimuli. To demonstrate the efficacy of PAWS for detecting hypersensitivity to noxious stimuli, we chemogenetically activated pain-aversion neurons in the amygdala, which further separated the behavioral representation of pain-related behaviors along a low-dimensional path. Taken together, this automated pain quantification approach should increase the ease and objectivity of collecting rigorous behavioral data, and it is compatible with other neural circuit dissection tools for determining the mouse pain state.