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Editorial article, editorial: neurological insights into communication and synchrony between others: what animal and human group communication can tell us.

research papers about animal communication

  • 1 Department of Psychology, University of Maryland, Baltimore, MD, United States
  • 2 Department of Biology and the Center for Translational Social Neuroscience, Emory University, Atlanta, GA, United States

Editorial on the Research Topic Neurological insights into communication and synchrony between others: what animal and human group communication can tell us

Communication is the cornerstone of human interaction, serving as the conduit through which ideas are exchanged, relationships are formed, and societies thrive. While we often think of communication using overt means, such as physical gestures and speaking to others, the intricacies of communication extend far beyond explicit communication, encompassing non-verbal cues and physiological actions that shape our understanding and interpretation of social interactions ( Phutela, 2015 ; Symons et al., 2016 ). In fact, given the recent advances in artificial intelligence, interpersonal interaction can be extended to occur not between individuals, but instead between individuals and computer, further complicating the role of covert cues. Additionally, recent advancements in neurological research in both animal and human studies have shed light on the underlying mechanisms of non-verbal behavior, offering profound insights into the complexities of human group dynamics and interpersonal communication (e.g., Hirsch et al., 2018 ). Indeed, understanding human communication may require delving into the methods and findings of animal models, as animal models have offered significant insight into human psychopathology ( Heller, 2016 ). Thus, in this Research Topic, we explore the intersection of human, artificial intelligence, and animal research regarding non-verbal communication. We highlight five seminal articles that contribute to our understanding of human communication utilizing diverse tools to understand these phenomena such as metacognition, animal models, and dyadic human interactions.

One article, Neurophysiological and Emotional Influences on Team Communication and Metacognitive Cyber Situational Awareness During a Cyber Engineering Exercise , demonstrates advancements in neurological technology and how they contribute to the understanding of human communication ( Ask et al. ). Researchers examine the realm of cyber operations, where human-to-human communication plays a pivotal role in achieving shared situational awareness for effective decision-making. Utilizing the Orient, Locate, Bridge (OLB) model, researchers investigate the neural correlates of metacognitive cyber situational awareness among cyber cadets. Their findings underscore the influence of neurophysiological and emotional factors on team communication, revealing the importance of vagal tone in shaping metacognitive judgments and mood. This study provides essential insights into the cognitive processes underlying effective communication in cyber defense, and more broadly, to hierarchical communication. Together, these results pave the way for innovative approaches to recruitment, education, and training in this critical domain.

A possible explanation for the relationship between vagal tone and communicative success in cyber operations is emotional state. Functional Graph Contrastive Learning of Hyperscanning EEG Reveals Emotional Contagion Evoked by Stereotype-Based Stressors shows how humans can transmit emotion to one another without being consciously aware ( Huang et al. ). Authors show how emotional contagion pervades dyadic interactions, shaping the dynamics of collaborative tasks and influencing performance outcomes. This article also employed EEG-based hyperscanning to unravel the neural mechanisms underlying emotional contagion in the context of stereotype-based stressors. Through functional graph contrastive learning (fGCL), researchers suggest the impact of emotional contagion on participants' neural activity patterns, revealing its substantial role in modulating performance trajectories. This study contributes valuable insights into the neural underpinnings of emotional dynamics in dyads, enriching our understanding of social interactions in diverse contexts.

Attention to not only one's emotional state, but one's physiological state, can impact communication and interpersonal synchrony, as shown in Autonomic Synchrony Induced by Hyperscanning Interoception During Interpersonal Synchronization Tasks ( Balconi et al. ). This article demonstrates that social interactions are inherently dynamic, characterized by reciprocal influences on emotional states and physiological rhythms. This work investigates the role of autonomic synchrony in dyadic interpersonal synchronization tasks, exploring the impact of interoceptive focus on physiological coherence. By employing hyperscanning techniques, researchers reveal higher synchrony between paired participants in heart rate variability (HRV), skin conductance level (SCL), and heart rate (HR) during tasks involving focused attention on one's own breathing. These findings highlight the interplay between interoception and interpersonal synchrony, offering new avenues for studying psychophysiological coherence in real-time social interactions. This research also shows how human communication and synchrony can be seen not only though explicit communication and neurological activity, but also cardiovascular responses.

Other research takes a more cellular approach, looking at mirror neuron systems (MNS; Bonini, 2017 ), which are essential in understanding the intentions and movements of others. In, Effects of Avatar Shape and Motion on Mirror Neuron System Activity , researchers explored the role of the MNS in perceiving humanness in avatars, shedding light on how avatar characteristics impact neural activity ( Miyamoto et al. ). Application of electroencephalogram (EEG) analysis demonstrated activation of the MNS in response to human-like avatar shapes and motions, highlighting the importance of considering both visual and kinematic cues in avatar design and interpreting the intentions of others through physical movement. These findings offer valuable insights for enhancing inter-avatar communication and fostering a sense of social presence in virtual environments. Further, they demonstrate how understanding human communication in humans is an automatic process that can extend beyond assessing other humans, even down to the neuronal level.

Animal research provides further evidence for the influence of non-verbal communication. In Listening to Your Partner: Serotonin Increases Male Responsiveness to Female Vocal Signals in Mice , researchers explore how the context surrounding vocal communication can significantly influence the perception of vocal signals ( Hood and Hurley ). Specifically, authors examined serotonin's role in modulating behavioral responses to vocal signals in mice. By manipulating serotonin levels systemically and locally in the inferior colliculus (IC), researchers uncover the nuanced effects of serotonin on vocal behavior, highlighting the neurotransmitter's role in modulating male responsiveness to female vocal signals. These findings underscore the importance of considering neurotransmitter systems in understanding the mechanisms of context-dependent communication.

In conclusion, the articles presented in this Research Topic offer a multifaceted exploration of non-verbal communication from neurological perspectives, spanning human and animal research domains. From cyber defense decision-making to avatar design in virtual environments, interpersonal synchrony in social interactions, emotional contagion in dyadic tasks, and autonomic synchrony to serotonergic modulation of vocal perception, these studies illuminate the diverse facets of non-verbal behavior and its underpinnings. By integrating insights from human and animal models, we can deepen our understanding of communication dynamics and pave the way for future advancements in understanding an innate human behavior.

Author contributions

RA: Conceptualization, Writing – original draft. MW: Conceptualization, Writing – review & editing.

The author(s) declare that no financial support was received for the research, authorship, and/or publication of this article.

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Publisher's note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

Bonini, L. (2017). The extended mirror neuron network: anatomy, origin, and functions. The Neuroscientist 23, 56–67. doi: 10.1177/1073858415626400

PubMed Abstract | Crossref Full Text | Google Scholar

Heller, A. S. (2016). Cortical-subcortical interactions in depression: from animal models to human psychopathology. Front. Syst. Neurosci. 10, 20. doi: 10.3389/fnsys.2016.00020

Hirsch, J., Adam Noah, J., Zhang, X., Dravida, S., and Ono, Y. (2018). A cross-brain neural mechanism for human-to-human verbal communication. Soc. Cogn. Affect. Neurosci. 13, 907–920. doi: 10.1093/scan/nsy070

Phutela, D. (2015). The importance of non-verbal communication. IUP J. Soft Skills 9, 43.

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Symons, A. E., El-Deredy, W., Schwartze, M., and Kotz, S. A. (2016). The functional role of neural oscillations in non-verbal emotional communication. Front. Hum. Neurosci. 10, 239. doi: 10.3389/fnhum.2016.00239

Keywords: synchrony, human communication, EEG, vocal signaling, non-verbal communication

Citation: Amey RC and Warren MR (2024) Editorial: Neurological insights into communication and synchrony between others: what animal and human group communication can tell us. Front. Hum. Neurosci. 18:1415166. doi: 10.3389/fnhum.2024.1415166

Received: 10 April 2024; Accepted: 15 April 2024; Published: 02 May 2024.

Edited and reviewed by: Lutz Jäncke , University of Zurich, Switzerland

Copyright © 2024 Amey and Warren. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY) . The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Rachel C. Amey, ramey.ameyc@gmail.com

This article is part of the Research Topic

Neurological Insights into Communication and Synchrony between others: What Animal and Human Group Communication can tell us.

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An Introduction to Animal Communication

research papers about animal communication

The ability to communicate effectively with other individuals plays a critical role in the lives of all animals. Whether we are examining how moths attract a mate, ground squirrels convey information about nearby predators, or chimpanzees maintain positions in a dominance hierarchy, communication systems are involved. Here, I provide a primer about the types of communication signals used by animals and the variety of functions they serve. Animal communication is classically defined as occurring when “...the action of or cue given by one organism [the sender] is perceived by and thus alters the probability pattern of behavior in another organism [the receiver] in a fashion adaptive to either one both of the participants” (Wilson 1975). While both a sender and receiver must be involved for communication to occur (Figure 1), in some cases only one player benefits from the interaction. For example, female Photuris fireflies manipulate smaller, male Photinus fireflies by mimicking the flash signals produced by Photinus females. When males investigate the signal, they are voraciously consumed by the larger firefly (Lloyd 1975; Figure 2). This is clearly a case where the sender benefits and the receiver does not. Alternatively, in the case of fringe-lipped bats, Trachops cirrhosus , and tungara frogs, Physalaemus pustulosus , the receiver is the only player that benefits from the interaction. Male tungara frogs produce advertisement calls to attract females to their location; while the signal is designed to be received by females, eavesdropping fringe-lipped bats also detect the calls, and use that information to locate and capture frogs (Ryan et al . 1982). Despite these examples, there are many cases in which both the sender and receiver benefit from exchanging information. Greater sage grouse nicely illustrate such “true communication”; during the mating season, males produce strutting displays that are energetically expensive, and females use this honest information about male quality to choose which individuals to mate with (Vehrencamp et al . 1989).

Figure 1 A model of animal communication.

Figure 2:  Photinus fireflies. Courtesy of Tom Eisner.

Signal Modalities

Animals use a variety of sensory channels, or signal modalities, for communication. Visual signals are very effective for animals that are active during the day. Some visual signals are permanent advertisements; for example, the bright red epaulets of male red-winged blackbirds, Agelaius phoeniceus, which are always displayed, are important for territory defense. When researchers experimentally blackened epaulets, males were subject to much higher rates of intrusion by other males (Smith 1972). Alternatively, some visual signals are actively produced by an individual only under appropriate conditions. Male green anoles, Anolis carolinensis, bob their head and extend a brightly colored throat fan (dewlap) when signaling territory ownership. Acoustic communication is also exceedingly abundant in nature, likely because sound can be adapted to a wide variety of environmental conditions and behavioral situations. Sounds can vary substantially in amplitude, duration, and frequency structure, all of which impact how far the sound will travel in the environment and how easily the receiver can localize the position of the sender. For example, many passerine birds emit pure-tone alarm calls that make localization difficult, while the same species produce more complex, broadband mate attraction songs that allow conspecifics to easily find the sender (Marler 1955). A particularly specialized form of acoustic communication is seen in microchiropteran bats and cetaceans that use high-frequency sounds to detect and localize prey. After sound emission, the returning echo is detected and processed, ultimately allowing the animal to build a picture of their surrounding environment and make very accurate assessments of prey location. Compared to visual and acoustic modalities, chemical signals travel much more slowly through the environment since they must diffuse from the point source of production. Yet, these signals can be transmitted over long distances and fade slowly once produced. In many moth species, females produce chemical cues and males follow the trail to the female’s location. Researchers attempted to tease apart the role of visual and chemical signaling in silkmoths, Bombyx mori , by giving males the choice between a female in a transparent airtight box and a piece of filter paper soaked in chemicals produced by a sexually receptive female. Invariably, males were drawn to the source of the chemical signal and did not respond to the sight of the isolated female (Schneider 1974; Figure 3). Chemical communication also plays a critical role in the lives of other animals, some of which have a specialized vomeronasal organ that is used exclusively to detect chemical cues. For example, male Asian elephants, Elaphus maximus , use the vomeronasal organ to process chemical cues in female’s urine and detect if she is sexually receptive (Rasmussen et al . 1982).

Figure 3 Male silkmoths are more strongly attracted to the pheromones produced by females (chemical signal) than the sight of a female in an airtight box (visual signal). Tactile signals, in which physical contact occurs between the sender and the receiver, can only be transmitted over very short distances. Tactile communication is often very important in building and maintaining relationship among social animals. For example, chimpanzees that regularly groom other individuals are rewarded with greater levels of cooperation and food sharing (de Waal 1989). For aquatic animals living in murky waters, electrical signaling is an ideal mode of communication. Several species of mormyrid fish produce species-specific electrical pulses, which are primarily used for locating prey via electrolocation, but also allow individuals searching for mates to distinguish conspecifics from heterospecifics. Foraging sharks have the ability to detect electrical signals using specialized electroreceptor cells in the head region, which are used for eavesdropping on the weak bioelectric fields of prey (von der Emde 1998).

Signal Functions

Some of the most extravagant communication signals play important roles in sexual advertisement and mate attraction. Successful reproduction requires identifying a mate of the appropriate species and sex, as well as assessing indicators of mate quality. Male satin bowerbirds, Ptilonorhynchus violaceus , use visual signals to attract females by building elaborate bowers decorated with brightly colored objects. When a female approaches the bower, the male produces an elaborate dance, which may or may not end with the female allowing the male to copulate with her (Borgia 1985). Males that do not produce such visual signals have little chance of securing a mate. While females are generally the choosy sex due to greater reproductive investment, there are species in which sexual roles are reversed and females produce signals to attract males. For example, in the deep-snouted pipefish, Syngnathus typhle , females that produce a temporary striped pattern during the mating period are more attractive to males than unornamented females (Berglund et al . 1997). Communication signals also play an important role in conflict resolution, including territory defense. When males are competing for access to females, the costs of engaging in physical combat can be very high; hence natural selection has favored the evolution of communication systems that allow males to honestly assess the fighting ability of their opponents without engaging in combat. Red deer, Cervus elaphus , exhibit such a complex signaling system. During the mating season, males strongly defend a group of females, yet fighting among males is relatively uncommon. Instead, males exchange signals indicative of fighting ability, including roaring and parallel walks. An altercation between two males most often escalates to a physical fight when individuals are closely matched in size, and the exchange of visual and acoustic signals is insufficient for determining which animal is most likely to win a fight (Clutton-Brock et al . 1979). Communication signals are often critical for allowing animals to relocate and accurately identify their own young. In species that produce altricial young, adults regularly leave their offspring at refugia, such as a nest, to forage and gather resources. Upon returning, adults must identify their own offspring, which can be especially difficult in highly colonial species. Brazilian free-tailed bats, Tadarida brasiliensis , form cave colonies containing millions of bats; when females leave the cave each night to forage, they place their pup in a crèche that contains thousands of other young. When females return to the roost, they face the challenge of locating their own pups among thousands of others. Researchers originally thought that such a discriminatory task was impossible, and that females simply fed any pups that approached them, yet further work revealed that females find and nurse their own pup 83% of the time (McCracken 1984, Balcombe 1990). Females are able to make such fantastic discriminations using a combination of spatial memory, acoustic signaling, and chemical signaling. Specifically, pups produce individually-distinct “isolation calls”, which the mother can recognize and detect from a moderate distance. Upon closer inspection of a pup, females use scent to further confirm the pup’s identity. Many animals rely heavily on communication systems to convey information about the environment to conspecifics, especially close relatives. A fantastic illustration comes from vervet monkeys, Chlorocebus pygerythrus , in which adults give alarm calls to warn colony members about the presence of a specific type of predator. This is especially valuable as it conveys the information needed to take appropriate actions given the characteristics of the predator (Figure 4). For example, emitting a “cough” call indicates the presence of an aerial predator, such as an eagle; colony members respond by seeking cover amongst vegetation on the ground (Seyfarth & Cheney 1980). Such an evasive reaction would not be appropriate if a terrestrial predator, such as a leopard, were approaching.

Figure 4 Vervet monkeys. Many animals have sophisticated communication signals for facilitating integration of individuals into a group and maintaining group cohesion. In group-living species that form dominance hierarchies, communication is critical for maintaining ameliorative relationships between dominants and subordinates. In chimpanzees, lower-ranking individuals produce submissive displays toward higher-ranking individuals, such as crouching and emitting “pant-grunt” vocalizations. In turn, dominants produce reconciliatory signals that are indicative of low aggression. Communication systems also are important for coordinating group movements. Contact calls, which inform individuals about the location of groupmates that are not in visual range, are used by a wide variety of birds and mammals. Overall, studying communication not only gives us insight into the inner worlds of animals, but also allows us to better answer important evolutionary questions. As an example, when two isolated populations exhibit divergence over time in the structure of signals use to attract mates, reproductive isolation can occur. This means that even if the populations converge again in the future, the distinct differences in critical communication signals may cause individuals to only select mates from their own population. For example, three species of lacewings that are closely related and look identical are actually reproductively isolated due to differences in the low-frequency songs produced by males; females respond much more readily to songs from their own species compared to songs from other species (Martinez, Wells & Henry 1992). A thorough understanding of animal communication systems can also be critical for making effective decisions about conservation of threatened and endangered species. As an example, recent research has focused on understanding how human-generated noise (from cars, trains, etc) can impact communication in a variety of animals (Rabin et al . 2003). As the field of animal communication continues to expand, we will learn more about information exchange in a wide variety of species and better understand the fantastic variety of signals we see animals produce in nature.

Vomeronasal organ – auxiliary olfactory organ that detects chemosensory cues

Altricial – the state of being born in an immature state and relying exclusively on parental care for survival during early development

Refugia – areas that provide concealment from predators and/or protection from harsh environmental conditions

Conspecifics – organisms of the same species

References and Recommended Reading

Balcombe, J.P. Vocal recognition of pups by mother Mexican free-tailed bats, Tadarida brasiliensis mexicana . Animal Behaviour 39 , 960-966 (1990). Berglund, J., Rosenqvist G. and Bernet P. Ornamentation predicts reproductive success in female pipefish. Behavioral Ecology and Sociobiology 40 , 145-150 (1997). Clutton-Brock, T., Albon S., Gibson S. & Guinness F. The logical stag: Adaptive aspects of fighing in the red deer. Animal Behaviour 27 , 211-225 (1979). de Waal F.B.M. Food sharing and reciprocal obligations among chimpanzees. Journal of Human Evolution 18 , 433–459 (1989).

Hauser, M. 1997. The Evolution of Communication . Cambridge, MA: MIT Press. Lloyd, J.E. Aggressive mimicry in Photuris: signal repertoires by femmes fatales. Science 197 , 452-453 (1975).

Marler, P. Characteristics of some animal calls. Nature 176 , 6-8 (1955). Martinez Well, M. & Henry C.S. The role of courtship songs in reproductive isolation among populations of green lacewings of the genus Chrysoperla . Evolution 46 , 31-43 (1992). McCracken, G.F. Communal nursing in Mexican free-tailed bat maternity colonies. Science 223 , 1090-1091(1984).

Rabin, L.A., McCowan B., Hooper S.L & Owings D.H. Anthropogenic noise and its effect on animal communication: an interface between comparative psychology and conservation biology. International Journal of Comparative Psychology 16 , 172-192 (2003). Ryan M.J., Tuttle M.D., & Rand A.S. Sexual advertisement and bat predation in a neotropical frog. American Naturalist 119 , 136–139 (1982). Schneider, D. The sex attractant receptors of moths. Scientific American 231 , 28-35 (1974). Seyfarth, R.M., Cheney D.L. & Marler P. Monkey responses to three different alarm calls: Evidence for predator classification and semantic communication. Science 210 , 801-803 (1980). Smith, D. The role of the epaulets in the red-winged blackbird, ( Agelaius phoeniceus ) social system. Behaviour 41 , 251-268 (1972).

Vehrencamp, S.L., Bradbury J.W., & Gibson R.M. The energetic cost of display in male sage grouse. Animal Behaviour 38 , 885-896 (1989). von der Emde, G. Electroreception. In D. H. Evans (ed.). The Physiology of Fishes , pp. 313-343. Boca Raton, FL: CRC Press (1998). Wilson, E.O. Sociobiology: The New Synthesis . Cambridge, MA: Harvard University Press (1975).

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Article Contents

1. introduction, 2. materials and methods, 3. analysis, 4. evaluation and discussion, 5. conclusion, acknowledgements, data availability, references–textbooks, the representation of animal communication and language evolution in introductory linguistics textbooks.

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Sławomir Wacewicz, Michael Pleyer, Aleksandra Szczepańska, Aleksandra Ewa Poniewierska, Przemysław Żywiczyński, The representation of animal communication and language evolution in introductory linguistics textbooks, Journal of Language Evolution , Volume 7, Issue 2, July 2022, Pages 147–165, https://doi.org/10.1093/jole/lzac010

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The last three decades have brought a wealth of new empirical data and methods that have transformed investigations of language evolution into a fast-growing field of scientific research. In this paper, we investigate how the results of this research are represented in the content of the most popular introductory linguistic textbooks. We carried out a comprehensive computer-assisted qualitative study, in which we inspected eighteen English-language textbooks for all content related to the evolutionary emergence of language and its uniqueness in nature, in order to evaluate its thematic scope, selection of topics, theories covered, researchers cited, structural soundness, currency, and factual accuracy. Overall, we found that the content of interest lacks a defined canonical representation across the textbooks. The coverage of animal communication was relatively broad, with some recurring classic examples, such as vervet monkeys or honeybees; this content was mostly structured around the ‘design features’ approach. In contrast, the coverage of topics related to language origins and evolution was much less extensive and systematic, and tended to include a relatively large the proportion of content of historical value (i.e. creation myths, ‘bow-wow’ theories). We conclude by making recommendations for future editions of textbooks, in particular, a better representation of important frameworks such as signalling theory, and of current research results in this fast-paced field.

Research on the origins and evolution of language origins has a troubled history of being viewed as unscientific (see e.g. Whitney 1873[1872 ], Doerfer 1973 , Fisiak 1985 ; for a discussion, see Kaplan 2021 ). However, as also stressed in a number of recent introductions to this field (e.g. Fitch 2010 , 2017 ; Gong et al. 2014 ; Dediu and de Boer 2016 ; Progovac 2019 ; Boeckx 2021 ), the field has transformed significantly. It no longer can be seen as armchair philosophizing producing just-so-stories ( Gould and Lewontin 1979 ; cf. Lewontin 1998 ), or a kind of ‘intellectual game’ ( Kendon 1991 : 202). Instead, it has progressed to regular research centred around the day-to-day addressing of solvable Kuhnian puzzles, including both empirical research and carefully constructed theoretical models. In particular, the perception of language evolution as a scientific field has been academically consolidated in terms of institutional indicators, such as prestigious flagship publications, research centres, funded projects, conferences, and journals (see esp. Dediu and de Boer 2016 ), to yield a program that is institutionally as well as theoretically progressive ( Lakatos 1978 ; for a discussion, see Żywiczyński 2018 : 200–202). In sum, it is fair to say that ‘[r]esearch on language evolution is undoubtedly among the fastest-growing topics in linguistics’ ( Nölle et al. 2020 ).

Has all of this progress also percolated to the teaching of linguistics, as reflected in the content of linguistics textbooks? Introductory textbooks are generally seen as places where Kuhnian ‘“normal science” is defined and acknowledged fact is represented’ ( Hyland 2004 : 105). They thus serve an important function of ‘canonizing discourse’ within a discipline ( Brown 1993 ; cf. Hyland 2004 ; Love 2006 ). That is, they provide an orthodox, coherent ‘epistemological map’ of the landscape of the discipline and what it is about ( Hyland 2004 : 105). In fact, many students view introductory textbooks as representing ‘concrete embodiments of the knowledge of their disciplines’ ( Hyland 2004 : 105). Textbooks are, therefore, an important source for students to develop academic literacy in a discipline and become acculturated into it ( Johns 1997 ; Love 2006 ). As Myers (1992 : 3) puts it, ‘writing and reading [textbooks] reproduces knowledge and reproduces academics’. As outlined above, language evolution research has reached the stage of Kuhnian ‘normal science’ and is playing an expanding and important role in the language sciences. As such, it should be well represented in introductory linguistic textbooks, given that these texts strive to offer a ‘balanced and uniformly excellent coverage of the full range of modern linguistics’ ( Fasold & Connor-Linton ), address ‘all the topics that a student will need in their study of language’ ( McGregor ), and ‘up-to-date coverage of all the important areas of linguistics’ ( O’Grady et al. ).

In 2016, in a conference abstract, we reported on an exploratory investigation of topics related to language evolution research in fourteen introductory-level linguistic textbooks, finding that language evolution was not given the same status as, for example, language acquisition, language change, language and the brain, language and culture, or language and society, which are most extensively covered. We also judged that ‘the teaching of language evolution to students of general linguistics rests on out-dated and largely inadequate conceptual frameworks, and fails to communicate major theoretical breakthroughs and empirical results’ ( Wacewicz et al. 2016 ). Here, we revisit this issue with more rigorous tools and an updated and extended scope. Specifically, we investigate what introductory textbooks say about two closely intertwined (see, in particular, the now-classic work by Hockett 1960 and Hauser et al. 2002 ) topics: the questions on how language emerged and how it differs from the communication of non-human animals (henceforth: animals). In this paper, we present a first-of-its-kind, large computer-assisted qualitative study, in which we inspected eighteen textbooks for all contents related to the evolutionary emergence of language and its uniqueness in nature, in order to evaluate its thematic scope, selection of topics, theories covered, researchers cited, structural soundness, currency, and factual accuracy. Our primary goal is to guide better-quality teaching to the new generations of linguists, but our study may also serve as a blueprint for similar projects in other areas relevant to linguistics, as well as in other disciplines.

2.1 Materials and design

To evaluate the popularity and influence of linguistic textbooks, we consulted Open Syllabus (OS), a non-profit service with a corpus of over nine million English-language syllabi from 140 countries ( https://opensyllabus.org ). 1 Based on the OS popularity metrics, OS appearances , and OS score (as of 20 January 2021—see Table 1 ), we selected sixteen general introductory textbooks to linguistics, which we complemented with two very recent textbooks. In our analyses, we worked with the most recent edition of each textbook available as of 20 January 2021.

The introductory linguistics textbooks analysed. Edition: most recent edition available as of 20 January 2021. Year: year of publication of the most recent edition; OS App, OS Score - appearances (App) and popularity score on a 1-100 scale (Score) indicators at Open Syllabus; Chapter (Animal Communication, Language Evolution): page count of the chapter dedicated to this category (if present), this number is inclusive of any sections in the chapter; Section: LTA - page count of the section dedicated to the subcategory Language Trained Animals (if present). Section: other - page count of the section dedicated to subcategories Animal Communication Behaviors and Animal vs Human Communication (if present).

2.1.1 Scope of the analysis

The content of interest for our analyses can be described as any content related to the origins of language and its status among animal communication systems, or more generally, a species-comparative or evolutionary perspective on language. This thematic range closely matches the scope of the Evolang , http://evolang.org/ , conference series, which is commonly regarded as the main conference in this field and often assumed as a reference point in language evolution research ( Bergmann and Dale 2016 ; Wacewicz and Żywiczyński 2017 ). Hence, in order to help operationalize the exact thematic profile of the content to be coded, we stipulated it as ‘content with a good fit to the Evolang conference’. Following up on Wacewicz et al. (2016 ; see also Bergmann and Dale 2016 , on the main Evolang topic clusters), we divided the content top-down into two main thematic categories of interest as Animal Communication and Language Evolution (comprising both explanations of how language originated and interdisciplinary research relevant to language evolution, but excluding purely historical language change). Including animal communication in addition to language evolution was motivated by the fact that the comparison of human and non-human communication systems represents one of the most central methods for the investigation of the evolution of language. Finding similarities and differences between aspects of human language and the communication systems of other animals can be informative as to the evolutionary foundations of language and evolutionary pressures shaping it. As such, the comparative methods are well represented in work on language evolution (e.g. Fitch 2010 ; Tallerman and Gibson 2012 ; Macmahon and McMahon 2013 ). For example, at the recent Joint Conference on Language Evolution ( https://sites.google.com/view/joint-conf-language-evolution/home ) in Kanazawa, Japan, in August 2022, out of the 196 plenaries, invited talks, talks, and posters, 43 (22%) were on animal communication and cognition. We coded passages appearing anywhere in the content of each book, including boxes, footnotes, captions, and exercises (but not indexes or references). Section 2.2 explains the details of the coding procedure, and Section 3.1 provides a more detailed breakdown and discussion of the final set of codes and categories.

2.2 Procedure

The material was coded by two expert coders (AP, AS) with the computer-assisted qualitative research software NVivo 1.3 ( QSR International Pty Ltd., released August 2020 ), in collaboration with two language evolution experts (SW, MP).

The cyclical coding/re-coding process was completed in five steps (cf. Saldaña 2015 ):

1) Training: Two coders were trained with a training set consisting of excerpts from two textbooks. The coders were instructed to identify and mark all passages related to language origins, language evolution, or animal communication as described above, assuming one sentence as a minimal passage and one paragraph as a maximum passage. Each of the identified passages was assigned a short label, that is, a code. The coders were instructed to adopt a bottom-up approach and use the topics discussed at the Evolang conferences as the reference point for coding. Two language evolution experts provided feedback.

2) Individual open coding: each textbook was open coded by the two expert coders, working independently of each other. The coders identified the relevant passages through a three-step procedure:

(a) The coder analysed the table of contents and the indexes of each textbook to pre-select the potentially most relevant chapters, sections, and pages for close reading;

(b) The coder then manually skimmed the full content of each textbook;

(c) Finally, the coder completed a series of targeted keyword searches for a broad range of general keywords (e.g. evolution, emergence, origins, etc.) and specific keywords including names of animal species, names of disciplines (e.g. archaeology), and key concepts (e.g. FOXP2).

The coders coded all identified passages in the bottom-up approach described above.

3) Consensus I: in the first consensus phase, the output files from both expert coders were merged into a single NVivo file, containing over 1000 codes. The coders discussed each coded passage to arrive at a consensual coding and synthesized the material into a hierarchical structure of codes and categories (cf. Saldaña 2015 ). The coding scheme was progressively updated in the process, resulting in a roughly twofold reduction in the number of codes.

4) Verification: in this step, two language evolution experts (MP, SW), working together, reviewed the correctness of the coding scheme. The two language evolution experts discussed each coded passage and recommended changes to be considered in the next consensus phase as discussion points. They also did an additional manual skimming of the full text of each book. Additionally, they evaluated the coded passages for the factual accuracy of the information contained therein. This resulted in an additional set of codes and their annotations (see below, Section 4.1), which were then incorporated into the NVivo codebook.

5) Consensus II: all passages marked during the verification phase as discussion points were discussed together by all four experts to determine consensual coding and consensual classification into categories. This led to a final total of 462 codes and coding categories, available in Supplementary Material .

3.1 Topics overview

The main categories of investigation are presented here in a top-down manner. First, we will deal with the question of how many textbooks mention our two main coding categories: Animal Communication and Language Evolution.

First, only the Language Files , O’Grady et al . 2 and Yule have chapters on Animal Communication, and only Yule has a chapter on Language Evolution. Five other textbooks have sections dedicated to Animal Communication. Four other textbooks have sections dedicated to Language Evolution (see Table 1 ). However, it has to be noted that these sections also differ in length, ranging from less than one page to several pages. Out of eighteen textbooks, three do not make any references to either Animal Communication or Language Evolution. For the remaining fifteen, references to Animal Communication and Language Evolution can be further sub-categorized (see Figs. 1 and 2 ). For Animal Communication, there were three emergent broad topics (subcategories), and for Language Evolution, there were two subcategories.

Numbers of references in the five main coding subcategories in each of the linguistics textbooks analysed.

Numbers of references in the five main coding subcategories in each of the linguistics textbooks analysed.

Proportion of the number of references in the two main categories (Animal Communication vs. Language Evolution) and their main subcategories.

Proportion of the number of references in the two main categories (Animal Communication vs. Language Evolution) and their main subcategories.

Main category : Animal Communication


Animal communication behaviours : References to specific animal communicative behaviours or general characterizations of aspects of animal communication,

Animal versus human communication : Comparisons of animal versus human communication,

Language-trained animals : References to research with language-trained animals. 3

Main category: Language Evolution

Language origin theories : References to explanations of how language originated. This category includes references to larger theoretical frameworks as well as more specific scenarios of language origins, most of them historical rather than contemporary. Examples include creation myths like the story of Babel, historical onomatopoeic hypotheses of language emergence, or modern-day scientific accounts such as Dunbar’s gossip theory of language origins ( Dunbar 1996 ).

Language evolution research : References to interdisciplinary research directly connected to the field of language evolution (see Section 2). This category includes passages that do not describe a particular account or scenario of language origins, but instead present results and data, mostly empirical and relatively recent, that provide an evidential basis for inferences about the evolutionary history of language, and empirical and theoretical building blocks for scenarios and theories. Examples include references to the relevant fossil record (e.g. hominin braincases or speech organs), or relevant genetic research (e.g. the FOXP2 gene in ancient humans). Importantly, other bodies of research that are often drawn on to inform evolutionary scenarios (such as on language ­acquisition, sign language, neurolinguistics, or historical linguistics 4 ) were not included as a default—we only did so when they were presented in a context relating them to the questions of the evolutionary emergence and development of language in human phylogeny. This decision is based on the fact that these areas of research on their own are not only independent subjects of linguistic study, but also because they are most frequently not discussed in an evolutionary framework or with respect to their potential implications for language evolution.

3.1.1 Results

There are two main general findings: firstly, references to Animal Communication (80.4% of coded references) are overall much more frequent than to Language Evolution (19.6%), and secondly, there is significant variation between textbooks in the extent of coverage and selection of topics.

Regarding the subcategories of Animal Communication, these are roughly distributed evenly within that category, with 35.5% of references to specific animal communicative behaviours, 32.7% of references comparing animal and human communication, and 32.8% of references discussing language-trained animals. Within the category Language Evolution, results are also evenly split, with 50.2% of references to language evolution research, and 49.8% of references to specific language origin theories.

Secondly, the number of references to Animal Communication and Language Evolution differs significantly from textbook to textbook. As already mentioned, three textbooks do not include references to these topics at all, and a number of them also only make very little reference to these topics. The bulk of references come from ten textbooks, which, however, still vary quite extensively in their coverage, from 32 ( Fasold & Connor-Linton ) to 318 references ( O’Grady et al ). An overview of the frequencies of topics by textbooks can be seen in Fig. 1 .

Moreover, as can be seen in Fig. 2 , textbooks also differ in how much they discuss a particular topic, for example, the degree to which they discuss animal versus human communication, language-trained animals, and the other categories.

3.2 Animal communication

3.2.1 animal communication behaviours.

Thirteen out of fifteen textbooks (87%, with the exception of Bauer and Hayes ) make references to specific animal communicative behaviours or general characterizations of aspects of animal communication. Among those, references to primate communication are most frequent (12 textbooks, 125 references), followed by birds (7 textbooks, 108 references), arthropods, especially bees (11 textbooks, 82 references), and mammals other than primates (12 textbooks, 56 references). Overall, textbooks differ quite strongly in which specific animal communicative behaviours they refer to. This also holds for references to species. Textbooks reference a total of sixty-three different species. However, forty-four (69.8%) of these species are only mentioned in one textbook, and only four species/clades (6.3%) are mentioned in five textbooks or more. These are vervet monkeys (five textbooks), chimpanzees (six textbooks), dogs (seven textbooks) and honeybees (eleven textbooks).

In addition, sometimes textbooks also make more general statements, for example, about monkeys or primates, fish, or songbirds. Some textbooks also mention more general properties of animal communication, for example, discussions of non-vocal communication and whether animal signals are innate or learnt. Furthermore, some textbooks also discuss the general properties of animal signs, such as animal signs as expressions of emotional states, graded instead of discrete signals, and their limited range of meanings.

3.2.2 Animal versus human communication

Fourteen out of fifteen textbooks (93.33%, with the exception of Dirven & Verspoor ) include an explicit comparison of human language with the communication systems of non-human animals. As outlined in Section 3.1, three of them have chapters dedicated to animal communication or chapters on animal versus human communication.

As shown in Fig. 3 , the two topics that are mentioned by most textbooks are human language as a species-specific trait (11 textbooks, 24 references), as well as discussions of animal versus human communication in the context of Hockett’s design features (11 textbooks, 183 references).

Topics in the subcategory ‘animal vs. human communication’ by the number of textbooks and references.

Topics in the subcategory ‘animal vs. human communication’ by the number of textbooks and references.

Another prominent topic is discussions of the biological differences between humans and animals. A number of textbooks also discuss animals’ ability to learn and understand human language and differences in the cognitive abilities of animals and humans. Also frequent are specific comparisons between humans and chimpanzees and the claimed distinction that human communication is stimulus-free whereas animal communication is stimulus bound.

The last group of topics is differences between the features of human language and animal communication that are not captured by Hockett’s design features. These include the degree of iconicity in human language versus animal communication, the pragmatic function of human language, differences in the mode of communication, the property of redundancy in communication systems, and turn-taking as a characteristic property of human communication.

In terms of the number of references to concepts, references to Hockettian design features of language far outweigh all other references.

For this reason, we are now going to turn to discussions of Hockettian design features of the language. Out of the eleven textbooks that mention such features, only four (36.4%) explicitly mention Hockett by name and six (55.55%) reference the characteristic properties of human language as ‘design features’. All eleven, however, mention properties that were popularized by Hockett, either directly using Hockett’s terminology or expressing the same concept (see Fig. 4 ).

Topics in the sub-subcategory ‘Hockettian design features’ by the number of textbooks and references.

Topics in the sub-subcategory ‘Hockettian design features’ by the number of textbooks and references.

3.2.3 Language-trained animals

Discussions of ‘language-trained animals’, that is, animals exposed to some kind of sign system, can also be found frequently in textbooks. They can be found in ten out of fifteen textbooks (66.67%, with the exception of Bauer , Dirven and Verspoor , Hayes , Hazen and Meyer ). Textbooks focus mostly on research with great apes, which are mentioned in ten textbooks, whereas studies with parrots and dogs are both mentioned in four textbooks (40%). Only one textbook also reports on research with dolphins.

Regarding the great apes, all ten textbooks mention language-trained chimpanzees, eight mention bonobos; seven mention gorillas, and two textbooks also mention language-trained orangutans.

A sceptical perspective on language-trained animals is prevalent with eight textbooks discussing scepticism towards these experiments, and seven textbooks discussing these experiments in the context of stimulus-response conditioning. Three textbooks explicitly discuss the question of to what degree language-trained animals can be said to have acquired symbols, and three textbooks discuss the question if chimpanzees have acquired syntactic knowledge. Five textbooks explicitly compare humans’ and primates’ abilities to learn a language.

3.3 Language evolution

References in the category Language Evolution can be found in fifteen textbooks. However, as already shown in Table 1 and Fig. 1 (cf. Section 3.1), textbooks differ quite significantly in how much space they give to this topic and also in which research and language origin scenarios they discuss. As already mentioned, only Yule has a chapter dedicated to language evolution, and only Denham & Lobeck , Finegan , Hayes , and McGregor have dedicated sections on language evolution. But even here the amount of space devoted to language evolution differs, ranging from less than one page to six pages. The other textbooks sometimes only discuss language evolution in passing.

3.3.1 Language evolution research

Regarding the topics discussed, twelve textbooks make reference to biological aspects of human evolution (fifty-three references). This includes references to the evolution and structure of the human brain (eight textbooks, thirteen references), the FOXP2 gene (seven textbooks, sixteen references), and the evolution and structure of the human speech organs (five textbooks, eight references). Other topics in this area that are also mentioned are the evolution of bipedalism (three textbooks, four references), changes of the digestive and respiratory system (one textbook, two references), and handedness (one textbook, one reference). One other frequent feature is that many textbooks (eight textbooks, thirteen references) discuss that there is still much scepticism towards language evolution research and that many proposals are speculative. Explicitly positive attitudes towards the field of language evolution and its future are only expressed in two textbooks ( Finegan and McGregor ). Other topics discussed in a number of textbooks are extinct hominins (six textbooks, fifteen references), archaeological evidence (five textbooks, six references), and that language evolution research is an interdisciplinary endeavour. An overview of the main topics discussed and their frequencies can be found in Fig. 5 .

Main topics in the subcategory ‘language evolution research’ by the number of textbooks and references.

Main topics in the subcategory ‘language evolution research’ by the number of textbooks and references.

As illustrated by Fig. 6 , textbooks also differ considerably in the potential timeframes of language emergence, with estimates in eight textbooks ranging from 50 000 years ago to as much as 500 000 years ago, and even older estimates for related capacities such as the evolution of ‘speech areas’ of the brain (2 million years ago) or the ‘capacity for language-type communication’ (2.6 million years ago).

Different timeframe estimations for the evolution of language by the numbers of textbooks and references. Some textbooks mention more than one estimate.

Different timeframe estimations for the evolution of language by the numbers of textbooks and references. Some textbooks mention more than one estimate.

3.3.2 Language origin theories

Twelve textbooks mention language origin scenarios (with the exception of Dirven & Verspoor , Hayes and Hazen ). As Fig. 7 shows, the most frequently mentioned factor is language as an evolutionary adaptation (seven textbooks, eleven references) and discussions of the evolution of the properties of the faculty of language (six textbooks, fourteen references). Notably, cultural evolution is only mentioned explicitly in one textbook ( McGregor , three references), which we consider a major oversight (see Section 4.3.2). Another frequently discussed topic is divine origins myths, which can be found in five textbooks (thirty-two references). Equally frequent are discussions of social factors involved in the emergence of language (five textbooks, thirteen references) and the question in which modality language began, that is, if it was primarily vocal, gestural, or multimodal at first (five textbooks, nine references). Other aspects of language origins scenarios are discussed infrequently. This includes references to onomatopoeic scenarios of language evolution (three textbooks, seven references), the role of tool use in language evolution (three textbooks, seven references), the relationship of music and language in language evolution (two textbooks, three references), and catastrophic origin theories (four textbooks, nine references). In contrast, explicit references to gradualistic theories of language evolution can only be found in one textbook ( Fromkin et al.). The same holds for the important concept of protolanguage as an intermediary stage before fully modern language (e.g. Tallerman 2012), which is only mentioned in one textbook ( Fasold & Connor-Linton ).

Topics in the subcategory ‘language origin theories’ by the number of textbooks and references.

Topics in the subcategory ‘language origin theories’ by the number of textbooks and references.

4.1. Suggestions and resources for improvement

In this section, we offer some short notes on suggestions for improvements for introductory textbooks, along with suggestions for good resources regarding language evolution research (see Box 1 ).

Fitch, W. T. (2010). The Evolution of Language. Cambridge University Press.

The best book-length introduction to the subject with a special emphasis on biological evolution and the evolution of speech.

McMahon, A., & McMahon, R. (2013). Evolutionary Linguistics . Cambridge University Press.

Part of the ‘Cambridge Textbooks in Linguistics’ series, the only introductory textbook on the subject so far.

Progovac, Liljana (2019). A Critical Introduction to Language Evolution: Current Controversies and Future Prospects . Springer.

Part of the ‘SpringerBriefs in Linguistics’ series, explicitly aimed at acquainting ‘scholars with recent developments outside their own research areas’. Contrasts ­sudden (saltationist) and gradualist approaches to language evolution and calls for an empirical research paradigm to study the co-evolutionary loop between language, brains, and genes.

Tallerman, M. & Gibson, K. R., eds. (2012). The Oxford Handbook of Language Evolution . Oxford University Press.

A comprehensive handbook on the subject, featuring sixty-five chapters written by international experts on all aspects of language evolution. It covers ‘insights from comparative animal behaviour’, ‘the biology of language evolution’, ‘the prehistory of language’, theories on the initial emergence of language and ‘language change, creation, and transmission in modern humans’.

Christiansen, M. H. & Chater, N. (2022). The Language Game: How Improvisation Created Language and Changed the World . Basic Books.

A recent monograph that focusses on the cultural evolution of language. It adopts a view of linguistic behaviour as a game of communicative charades played over multiple generations. This process in turn explains the emergence of language and its structure.

Planer, R.J. & Sterelny, K. (2021). From Signal to Symbol: The Evolution of Language . MIT Press.

A novel, integrative account of language evolution, treating language as a result of successively developing protolanguages over the last two million year. The book focusses on archaeological data and the social and sociocognitive infrastructure enabling the emergence of language.

Hurford, J. R. (2014). Origins of language: A slim guide . Oxford University Press.

A condensed introductory version of Hurford’s two-part opus magnum on ‘Language in the Light of Evolution’ (2007, 2012) that focusses on the origins of meaning and the origins of grammar from an interdisciplinary perspective.

Fitch, W. T. 2017 (eds.) Special Issue on the Biology and Evolution of Language. Psychonomic Bulletin & Review 24 .

A collection of thirty-six Articles by international experts focussing on different empirical perspectives on language evolution from evolutionary biology, neuroscience, palaeoanthropology, comparative psychology, cultural evolution, linguistics and cognitive science.

Engesser, S. & Townsend, S. W. (2019). Combinatoriality in the vocal systems of nonhuman animals. WIREs Cognitive Science 10(4), e1493.

An article that reviews the current evidence for combinatorial systems in the vocal communication of different animals.

Tamariz, M. (2017). Experimental studies on the cultural evolution of language. Annual Review of Linguistics , 3, 389-407.

Gives an overview of experiments on cultural language evolution, emphasizing the role interaction and transmission play in the emergence of linguistic structure.

Krebs, J. R. & Dawkins, R. (1984). “Animal Signals: Mind-Reading and Manipulation”, in Behavioral Ecology: An Evolutionary Approach , eds. J. R. Krebs and R. Dawkins (Oxford: Blackwell), 380–402.

This groundbreaking work laid the foundation for signalling theory. It argues that the cooperative design of human language that we naturally take for granted is an exception rather than the rule in animal communication, and requires special conditions to emerge.

Overall, we found that the number of problematic passages about language evolution and animal communication was relatively low, with a total of sixty problematic passages of different dimensions found in all textbooks (thirty-eight for animal communication and twenty-two for language evolution). 5 The most frequent inaccurate statement concerns non-human primate vocal anatomy, with the claim that the vocal tracts of non-human primates are anatomically incapable of producing speech sounds. This has been demonstrated by more recent research to be inaccurate (e.g. Fitch 2000 ; Fitch et al. 2016 ). However, given that this is a relatively recent research development, it is understandable why it hasn’t permeated introductory textbooks yet.

One aspect we judged to be more problematic is that some textbooks ( Akmajian et al., Fasold & Connor-Linton , Rowe & Levine) make extensive reference to popular press publications, which in addition sometimes are also quite outdated. For example, in the 2017 edition of their textbook, Akmajian et al. cite a 1989 National Geographic Magazine article on the speech capacities of Neanderthals.

In addition, some textbooks, including Yule , cite and draw conclusions from research on the famous alarm call system of vervet monkeys alone (e.g. Cheney and Seyfarth 1990 ). Here, we suggest that introductory textbooks could profit from including more recent work on more complex signalling systems, such as those found in putty-nosed monkeys ( Arnold and Zuberbühler 2006 ) and Campbell’s monkeys ( Zuberbühler 2002 ; Ouattara et al. 2009 ), among others (see e.g. Townsend and Manser 2013 ; Engesser and Townsend 2019 ; Suzuki and Zuberbühler 2019 , for recent overviews).

In Box 1 , we list some useful recent resources that introductory textbooks could use to improve their representation of language evolution research (see also Box 2 ). In addition, Johansson (2020) offers a comprehensive bibliography of language evolution research.

There are a number of important alternatives to the design features framework that do not receive enough attention in textbooks. In particular, the distinction into the Faculty of Language in the broad versus narrow sense (FLB vs. FLN) , influentially put forward 20 years ago by Chomsky and collaborators ( Hauser et al. 2002 ; Fitch et al. 2005), is only briefly discussed in one textbook. Notwithstanding its serious limitations (see e.g. Wacewicz et al. 2020 ), the FLN/FLB distinction is now well established in the study of language.

Equally important are the approaches to both the evolution and uniqueness of language that highlight its social grounding and cognitive-interactional aspects (esp. Tomasello 1999 , 2008 ; Levinson 2006 ; Levinson and Holler 2014 ). Such approaches underscore the centrality of the cooperative nature of language as well as the cognitive infrastructure that enables language; the former completely eludes the Hockett-type classifications, and the latter is only indirectly or marginally present in features such as displacement.

Even if no particular theoretical formulation has yet reached textbook status, a forming consensus in the language evolution community is that the evolutionary history of cooperation in the hominin line plays a central role in the problem of language emergence ( Zlatev 2014 ). Cooperation is understood here widely and inclusive not only of the Gricean notion of cooperation but also signalling theory with its focus on cooperation as information donation (discussed in Section 4.2.2).

With regard to the cognitive infrastructure necessary for language , current research in language evolution focuses on prerequisites for communication based on the attribution of intentions ( Levinson 2006 ), Theory of Mind and social motivations ( Tomasello 1999 , 2008 ; Botha 2020 ), ostensive signals ( Scott-Phillips 2014 ), bodily mimesis ( Zlatev 2014 ) as well as turn-taking ( Levinson 2006 ; Levinson and Holler 2014 ).

4.2. Animal communication

4.2.1 hockett’s design features.

Section 3.2.2 reveals that the received way of comparing language with animal communication is through the list of ‘design features’ based more or less closely on those proposed by Hockett (esp. 1960 , but see also Hockett 1958 , 1959 ; 1966 ). This is understandable, given that the Hockettian system is still the classic, best known, and widely accepted system of comparing animal communication to language (e.g. Beecher 2021 ) and has been highly influential both in and beyond linguistics, in fields such as ethology (e.g. Hinde 1982 ), biosemiotics ( Noth 1990 ), and also language evolution (e.g. Fitch 2010 ; Cuskley 2020 ). However, more than 60 years have passed since Hockett’s first proposals ( 1958 , 1959 ), meaning that some of the deeper theoretical foundations of Hockett’s system rest on an outdated understanding of both biology and linguistics.

Biologically, Hockett’s approach is implicitly informed by once-popular misconceptions including that of evolution proceeding exclusively through incremental progressions, or anthropocentric views of animal communication systems as ‘incomplete languages’. We suggest that while Hockett’s design features might prove a good entry point to comparisons of human language and animal communication systems, textbooks would profit from extending their discussions by including insights from signalling theory (see Section 4.2.2). In contrast, Hockett’s approach rests implicitly on the ‘classic ethological’ model of communication, which is no longer supported in behavioural ecology (see Searcy and Nowicki 2005 : 7–9). From the point of view of linguistic theory, the Hockettian system inevitably reflects the mid-20 th -century understanding of what language is, with a heavy focus on its formal and structural properties; even if not incorrect, this understanding is at least incomplete, through its neglect of more recent cognitive, functional and interactionist perspectives (see esp. Wacewicz and Zywiczynski 2015 ).

In sum, the Hockettian system of design features has long been a cornerstone of understanding the status and origins of language, and its classic status still translates into its considerable descriptive and educational utility. However, textbooks would benefit from an overhaul of this system to the inclusion of more recent perspectives. Small steps in this direction are already evident in some textbooks. For example, O’Grady et al . feature a text box on ‘Updating the Design Features of Language’ with reference to a recent paper by Hauser et al. (2014) . However, these attempts are unsystematic and not yet standardized. In Box 2 , we briefly describe several points that we see as deserving inclusion.

4.2.2 Signalling theory

In biology, the received approach to studying the form, function and in particular evolution of animal communication is signalling theory: it is ‘the main body of theory applied to animal communication’ ( Power 2014 : 50), and it underlies contemporary textbooks on this topic (e.g. Maynard Smith and Harper 2003 ; Searcy and Nowicki 2005 ; Bradbury and Vehrencamp 2011 ). Signalling theory is a theoretical framework that applies neo-Darwinian principles to the study of communication. From that perspective, it construes communicators as agents designed to maximise their evolutionary fitness, and communication, like all behaviour, is seen as a tool for such fitness maximization. This perspective is interested in the underlying economics of communication, that is, what the animal stands to lose versus what it stands to gain from the communicative interaction. These costs and benefits ultimately affect the animal’s evolutionary fitness, and in this way, they translate into selection pressures shaping the evolution of communication systems. In particular, a central tenet of signalling theory is that the ‘goal’ of communication, understood from that perspective as its evolved function, is not to provide information to others—since making investments into the fitness of unrelated individuals instead of one’s own would be biologically inexplicable. Rather, the ‘goal’ of communication is to pursue one’s own fitness-enhancing goals, such as, for example, advertising one’s biological quality to potential mates, which can be trivialized into saying that communication is ‘self-interested’.

Signalling theory thus elucidates the deeper design principles of all animal communication. It is, therefore, fundamental from the species-comparative perspective, as it provides the only established unified framework to study all animal communication systems that do not presuppose language as a special case or a reference point. Signalling theory makes it possible to understand the design reasons for the limited range of expression in non-human animals (esp. Heintz and Scott-Phillips 2022 ), or explain certain unexpected patterns in the relation between communicative and cognitive capacities, such as the advanced communication system in honeybees (e.g. Wacewicz and Żywiczyński 2018 ). Most importantly, this perspective also clearly and independently identifies important ways in which human language stands out in nature (cf. esp. Knight 2016 ; Dessalles 2020 ). This is mostly related to the through-and-through cooperativeness of language, which is taken for granted in linguistics, but not normally expected in animal communication systems. This is as important for the species-comparative perspective as it is for research on the evolutionary origins of language, where the emergence of cooperative signalling as well as its continued evolutionary stability is considered a ‘central problem’ (e.g. Maynard Smith and Harper 2003 ; Fitch 2010 ; Heintz and Scott-Phillips 2022 ). Adding the perspective of signalling theory to textbooks would improve the theoretical context for discussing communication systems, and the differences and similarities between human language and non-human animal communication systems.

4.3 Language origins

Sections 3.3.1 and 3.3.2 reveal a relative paucity of information on those topics, especially when compared to much more extensive and detailed coverage of animal communication (see Fig. 2 ). While a majority of textbooks include mentions of language origins and language evolution research, only one textbook has a chapter on this topic and two more have sections extending beyond a single page. Further, we see a disproportionate representation of content of purely historical value, in particular, divine or mythical accounts of language origins (such as the story of Babel), or accounts developed in the 19th century (e.g. the ‘bow-wow’ or ‘pooh-pooh’ theories of language emerging from mimicking animals or from emotional cries). The inclusion of such themes may be substantiated on historical grounds as classic points of reference in the debates of language origins; however, their over-representation relative to the current state of the art may be felt as painting an unfavourable picture of contemporary language evolution research.

A particularly interesting example is Yule , the only textbook with a whole chapter dedicated to the topic of language origins. It is divided into sub-chapters representing several ‘sources’ of language: the divine source, the social interaction source, the natural sound source, the tool-making source, the physical adaptation source, and the genetic source. This presentation is problematic on a number of grounds. First, such a structuring juxtaposes explanations on incomparable levels ( Tinbergen 1963 ): implementation, versus adaptive function, versus phylogeny—that is the underlying physiological implementation of a feature, versus the selection pressures shaping it, versus its development in the species. For example, the level of physical implementation, here discussed in terms of the speech-adapted anatomy of the vocal tract, is listed in parallel to the level of genetic specification, even though the former is clearly causally dependent on the latter rather than being a ‘different source’. Furthermore, a juxtaposition of natural and supernatural ‘sources’ poses the danger of casting unscientific approaches as equal-status contenders—not only through the structure of subchapters already mentioned, but occasionally through unfortunate phrasing, for example, ‘If human language did emanate from a divine source, we have no way of reconstructing that original language, especially given the events in a place called Babel, <because the Lord did there confound the language of all the earth,> as described in Genesis’ (11: 9).

Another notable example is McGregor . This textbook presents an explicitly favourable evaluation of the field of language evolution research (pages 265, 267—the only such textbook except for Finegan ), and it offers a relatively brief but accurate and up-to-date coverage of this field, with a good selection of the most influential theories and relevant bodies of evidence. On the other hand, McGregor inexplicably extends Max Müller’s derogatory ‘bow-wow’ terminology to these approaches, using the labels ‘noddy’ (the gestural approach), ‘yackety-yack’ (Dunbar’s grooming hypothesis), and the ‘just genes’ theory, further subdivided into the ‘oops!’ and ‘chatting-up’ theories, which stand for the views of Chomsky and Pinker, respectively. As noted, for example, by Sprinker (1980 : 117), ‘it is hard to suppress one’s natural amusement when discussing them under such labels. […] [I]t is sometimes difficult for us to [take such ideas seriously], particularly with the persistent ring of Muller’s appellatives in our ears’.

4.4 Language evolution research

4.4.1 genetic, fossil, and material-culture data sources for modern language evolution research.

As shown in Section 3, textbooks do not rely extensively on the bodies of data that go beyond comparative evidence from animals. Of these, genetic evidence, and specifically the FOXP2 gene, is referenced relatively frequently (seven books, sixteen references), but information provided on this particular topic is often problematic (see Appendix A.1 in Supplementary Material). Palaeoanthropological and archaeological information is typically absent from textbooks or only mentioned as isolated facts ( fossil record : three books, four references; extinct hominins : six books, fifteen references; archaeology : five books, six references).

Still, the recently available genetic, fossil, and material-culture record of extinct hominins does allow us to shed considerable light on their linguistic abilities. Arguably the most interesting example, also for its significance to the science of language, is the strong interdisciplinary evidence pointing to the presence of recognizably human-like forms of communication in Neanderthals (see esp. Botha 2020 , for a critical overview). First cases for Neanderthal language backed up by comprehensive multidisciplinary evidence became available already a decade ago (e.g. Dediu and Levinson 2013 ), and additional evidence has accrued since then, such as on systematic neanderthalensis—sapiens interbreeding, the presence of the same derived mutations of the FOXP2 gene, very similar speech production and hearing apparatuses, and behavioural modernity of Neanderthals (see e.g. Johansson 2015 , Dediu and Levinson 2018 , for reviews).

Of course, we cannot say with absolute certainty whether Neanderthals did or did not have language ( Berwick et al. 2013 ; Botha 2020 ). Nevertheless, the perception that arguments regarding language in the prehistoric past must solely rely on speculation is no longer true. The last two to three decades have dramatically expanded the evidential basis for inferences about the language capacities of ancient hominins, making such inferences more robust through grounding them in multiple lines of converging evidence. Further, as shown above, it currently appears that the extent and weight of this converging evidence make it reasonable to change the null hypothesis: the bulk of evidence makes assuming the presence of language in Neanderthals more parsimonious than assuming its absence. Finally, this case is arguably of global importance to linguistics at large, as it is one of the very few potential ‘windows’ on dating the beginnings of language. Linguistic literature, including introductory textbooks (see Section 3.3.1) will keep on making estimates of the anciency of language, speculative as they are. We believe that basing these estimates on state-of-the-art information is no less important than hedging them for speculativeness.

4.4.2 Cultural evolution

The second cluster of empirical results relevant to language evolution comes from modelling and experimental studies focussing on the cultural evolution of language (e.g. Kirby 2012 ; Tamariz and Kirby 2016 ; Tamariz 2017 ). 6 This research is typically less concerned with the questions of the biological origin of the language faculty, but instead elucidates the emergence of meaning and structure in the linguistic code itself (see Section 4.3.2). This type of research now constitutes the core of empirical studies presented at Evolang conferences, but is virtually absent from linguistics textbooks.

Cultural evolution as a framework is particularly valuable, since it provides the most direct data on language evolution, and in particular on the evolutionary pressures shaping language structure. While comparative evidence from fields such as primatology or palaeoanthropological evidence from the hominin fossil record is necessarily indirect, the cultural evolution of communication can be studied in the laboratory, through experiments with human participants. As such, it represents a quintessentially hypothesis-testing approach, and the adoption of such methods to study the evolution of language was in a large part responsible for the change of profile of language evolution as a research field, from speculative to evidence and experiment based.

In stark contrast to its central role in current language evolution research, barring a single exception ( McGregor , three references), the topic of the cultural evolution of language is absent from the introductory linguistic textbooks that we examined. This can be partly explained by the relative recency of cultural evolution as a research area (e.g. Richerson and Boyd 2005 ; Mesoudi et al. 2006 ), both in a general sense and with regard to its role in language origins and language change. However, from today’s perspective, the almost complete neglect of this topic in introductory linguistic textbooks should be seen as a major oversight. Cultural evolution deserves a better representation in textbooks, as it addresses many of the concerns that language evolution research lacks an experimental, predictive framework. Furthermore, it is not only relevant for the study of the evolutionary origins of language, but provides a set of experimental methods that can be used to study the emergence of semiotic systems (e.g. Galantucci et al. 2012 , see Section 4.3.1). In addition, it can also be used to study the cultural change of languages more generally from an interdisciplinary perspective, in essence representing an analogue to the biologist’s Drosophila (cf. Roberts 2017 ). Integrating these research frameworks into discussions of language evolution in introductory textbooks is therefore likely to prove highly profitable.

In sum, in our examination of the most popular ‘introduction to linguistics’ textbooks, we found that the topic of language evolution and language origins is not prominently represented. Among the research areas that were most frequently mentioned were biological aspects of human evolution, references to extinct hominins, and discussions of the scepticism towards language evolution research. While animal communication is discussed much more frequently, textbooks also show wide variation in how much space they devote to the topic. Some of the most extensively covered topics include animal communicative behaviours, comparisons of animal versus human communication, and references to research with language-trained animals.

We found the factual accuracy to be good overall, with the most frequent mistake relating to the long-held belief that the vocal tract of non-human primates is incapable of producing speech sounds, which is understandable given that this is a relatively recent research development. However, the topics covered in our analysis are treated as a peripheral rather than core linguistic topic. We found the content on animal communication and in particular language origins and evolution to vary considerably between the textbooks, and this missing homogeneity is indicative of a lack of consensus on a canonical body of knowledge that students of linguistics should be expected to internalize. They also appear to be considered marginal, ‘luxury’ topics that are sometimes not covered at all or just in passing, especially in the shorter textbooks. An indicative example of this can be seen in O’Grady et al ., where animal communication is relegated to an online-only chapter only accessible via subscription (cf. Section 3.1).

The most important conclusion of our study is that the representation of the topics covered in our analysis, and in particular the origins and evolution of language, does not reflect the state of the art. Furthermore, this problem appears to disproportionately concern the most influential textbooks as measured by their OS score (esp. Fromkin , Yule— see Section 1 and Table 1 ), whose consecutive editions to a large degree inherit their content and organization from previous editions, which favours only local instead of global changes.

We recommended amendments on several dimensions. First, the relatively extensive representation of non-scientific content such as myths, religious stories, and 19th-century ‘bow-wow’-type accounts to the detriment of presentations of modern language evolution research, which was perhaps merited several decades ago, is no longer tenable in the face of the wealth of currently available scientific content that informs language origins questions. Non-scientific motifs should be reduced to symbolic proportions that can be justified on the grounds of their historical importance as cultural codes, in favour of a better representation of contemporary accounts, debates, and evidence (see esp. Section 4.3.1). Second, several specific approaches or perspectives that are at the heart of current language evolution research are absent from linguistics textbooks, in particular signalling theory (Section 4.2.2) and cultural evolution (Section 4.3.2). Their inclusion would be highly beneficial to young adepts of linguistics, for whom it would be an early (and in many cases perhaps the only) opportunity to become aware of dynamically developing approaches to the study of language-related phenomena. Third, although abandoning the reliance on Hockett-type lists of design features would be neither feasible nor fully desirable, such lists should be redefined to accommodate the recent findings such as on the status of arbitrariness in language or the importance of concepts such as cooperativeness, domain generality and turn-taking (Sections 4.2 and 4.2.1).

One limitation of this study is that we only examined English-language textbooks published which, with the exception of Dirven and Verspoor (2004) , were published in the USA and UK. For future studies, it would be interesting to see if this pattern is also apparent in introductory linguistics textbooks in other languages, which might come from different disciplinary and intellectual traditions.

We are grateful to Arkadiusz Jasiński, who helped develop an early version of this study. We would also like to thank the two anonymous reviewers as well as Dan Dediu for their insightful comments on the manuscript.

This research was supported by the Polish National Science Centre under grant agreement UMO-2019/34/E/HS2/00248. MP was supported by a postdoctoral fellowship from the University Centre of Excellence IMSErt: Interacting Minds, Societies, Environments, Nicolaus Copernicus University in Toruń.

Conflict of interest statement : The authors declare that there is no conflict of interest regarding the publication of this article.

The data underlying this article are available in OSF, at https://osf.io/5ta7x/

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It is of course important to note that this selection of textbooks is biased in that it is limited to textbooks in English, which come with their own biases in terms of a predominant focus on English as the language discussed most frequently. This bias in the Open Syllabus list is very likely due to the fact that although it covers 140 countries in total, it is a list of English-language syllabi and the majority of syllabi are taken from schools in English-speaking countries. In a future study, it would be highly interesting if the patterns found in this paper regarding English-language linguistics textbooks also hold for textbooks in other languages.

However, in O’Grady et al. , the chapter on animal communication is an ‘online only’ chapter that is only accessible with a subscription to the Macmillan Launchpad Solo service.

Language-trained animals were chosen as a subcategory on the basis of their frequent discussion both in linguistics textbooks and language evolution textbooks with relation to language evolution, and because the capacities of language- and symbol-trained animals have been of central interest to researchers in language evolution for a very long time (e.g. Lyn 2012 ).

While it could be argued that the history of languages is directly relevant to language evolution, most of the textbooks did not make explicit references to language evolution when discussing historical language change. This is in line with the widely held view in traditional historical linguistics that a ‘topic not generally considered to be properly part of historical linguistics is the ultimate origin of human language and how it may have evolved from non-human primate call systems, gestures, or whatever, to have the properties we now associate with human languages in general’ ( Campbell 2013 : 2; see Hartmann 2020 , for discussion).

For a comprehension description of problematic passages, see the Appendix A.1 in Supplementary Data.

For a short description of research on the cultural evolution of language, see the Appendix A.2 in Supplementary Data.

Supplementary data

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AP®︎/College Biology

Course: ap®︎/college biology   >   unit 8.

  • Intro to animal behavior
  • Innate behaviors
  • Learned behaviors

Animal communication

  • Animal behavior: foraging
  • Responses to the environment
  • Communication is when one animal transmits information to another animal causing some kind of change in the animal that gets the information.
  • Communication is usually between animals of a single species, but it can also happen between two animals of different species.
  • Animals communicate using signals , which can include visual; auditory, or sound-based; chemical, involving pheromones ; or tactile, touch-based, cues.
  • Communication behaviors can help animals find mates, establish dominance, defend territory, coordinate group behavior, and care for young.


Communication takes many forms.

  • Pheromones—chemicals
  • Auditory cues—sounds
  • Visual cues
  • Tactile cues—touch

Auditory signals

  • Monkeys cry out a warning when a predator is near, giving the other members of the troop a chance to escape. Vervet monkeys even have different calls to indicate different predators.
  • Bullfrogs croak to attract female frogs as mates. In some frog species, the sounds can be heard up to a mile away!
  • Gibbons use calls to mark their territory, keeping potential competitors away. A paired male and female, and even their offspring, may make the calls together.

Visual signals

Tactile signals—touch, what is communication used for.

  • Obtaining mates. Many animals have elaborate communication behaviors surrounding mating, which may involve attracting a mate or competing with other potential suitors for access to mates. See more information. Communication behaviors surrounding mating are often highly ritualized. For instance, a male may perform an intricate dance, show off decorative features—such as bright patches or elaborate patterns—or perform a characteristic song to attract a female. Similarly, males may compete with each for mates other using ritualized display behaviors , which usually involve posturing and gestural or vocal "threats" rather than actual fighting.
  • Establishing dominance or defending territory. In many species, communication behaviors are important in establishing dominance in a social hierarchy or defending territory. See more information. Communication, for example, may allow disputes over status or territory to be settled without the need for fighting. By posturing, vocalizing, or making aggressive gestures, both participants make a relatively honest advertisement of their ability and willingness to fight. This allows both parties to size each other up, and the weaker may voluntarily back down.
  • Coordinating group behaviors. In social species, communication is key in coordinating the activities of the group, such as food acquisition and defense, and in maintaining group cohesion. See more information. Communication may be used, for example, to direct other group members to a food source. Honeybee foragers use the waggle dance for this purpose, and ants use pheromone trails. Pack-hunting predators, such as wolves, also communicate to capture prey as a group. Group members may signal to coordinate defensive behaviors. For example, this is the case when a crushed ant incites other ants to swarm, or when a monkey gives an alarm call upon spotting a predator. Communication behaviors can also maintain cohesion within a group or establish social bonds and relationships. For instance, grooming among primates fosters cooperation and cohesion among group members.
  • Caring for young. Among species that provide parental care to offspring, communication coordinates parent and offspring behaviors to help ensure that the offspring will survive. See more information. Tactile signals exchanged between newborn animals and their mothers, for example, trigger the mother to provide food and may also stimulate the formation of parent-child bonds through hormone release. Gull chicks tapping on the red spots on their parents' beaks—see article on innate behavior —is another example of a communication behavior that favors the survival of offspring.


  • " Behavioral biology: Proximate and ultimate causes of behavior " by OpenStax College, Biology, CC BY 4.0 ; download the original article for free at http://cnx.org/contents/[email protected] .
  • " Communication behavior in animals " by Douglas Wilkin and Jean Brainard, CC BY-NC 3.0

Works cited

  • Eric Gillam, "An Introduction to Animal Communication," Nature Education Knowledge 3, no. 10 (2011): 70, http://www.nature.com/scitable/knowledge/library/an-introduction-to-animal-communication-23648715 .
  • Peter Tyson, "Dogs' Dazzling Sense of Smell," Nova, last modified October 4, 2012, http://www.pbs.org/wgbh/nova/nature/dogs-sense-of-smell.html .
  • Duncan E. Jackson and Francis L. W. Ratnieks, "Communication in Ants," Current Biology 16, no. 15 (2006): R570-R574, http://dx.doi.org/10.1016/j.cub.2006.07.015 .
  • "Trail Pheromone," Wikipedia, last modified April 11, 2016, https://en.wikipedia.org/wiki/Trail_pheromone .
  • "Ant," Wikipedia, last modified June 18, 2016, https://en.wikipedia.org/wiki/Ant .
  • "Chemical Pheromone Communication Between Ants," antARK, accessed June 18, 2016, https://antark.net/ant-life/ant-communication/ant-pheromones/ .
  • "Dog Communication," Wikipedia, last modified June 17, 2016, https://en.wikipedia.org/wiki/Dog_communication .
  • "Dog Behavior," Wikipedia, last modified June 9, 2016, https://en.wikipedia.org/wiki/Dog_behavior .
  • "Whale Vocalization," Wikipedia, last modified June 8, 2016, https://en.wikipedia.org/wiki/Whale_vocalization .
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10.2: Human Language versus Animal Communication

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Human Language versus Animal Communication, from Sarah Harmon

Video Script

part of it. I know a number of you have felt it was difficult throughout this whole process; I’m not going to argue. But when we talk about the actual neural components to language—how the brain processes language—this is where things get really technical.

For this section on animal vs human communication, this is evidence that we have up to this point. It's still evolving and we're still learning, but I can be more confident with what we're saying and what we're presenting here. There is so much that we're still learning about this thing between our ears and how it processes everything, including language, that this is constantly evolving. Just for a frame of reference, I am recording this mid-August of 2021; by the time you watch this, there could be some significant advances, and I won't know it until we get there. Just know that everything I’m presenting is with a grain of salt, in the sense of I am presenting the latest that we have up to this point. There is much more research to come, there is also more research underway and, additionally, our understanding of everything is changing. I'm confident and what I’m going to present for this chapter, but understand that things are changing.

Let's start off with a discussion that we had back in Chapter 1 when we talked about animal communication and human language. Let's refresh our memories a bit; let's go back to those attributes or hallmarks of human language . We understand that human beings have something that is potentially unique, if not very rare, that this form of communication—we talk with arbitrary signs and signals, we talk about things in front of us and not in front of us., we transmit aspects of our culture of our lives, we make an infinite use of finite means, we only have a certain number of vocabulary or lexicon, we only have a certain number of ways that we can put them into a phrase, and yet we can say anything that comes to mind, that we are productive and creative and constantly changing and adapting with our language, we can talk about all these things and ourselves, we can create, we can express and we can talk about things that are not just here in now, not just immediate needs, we go well beyond that. Just think of this class alone!

There are some similar aspects that we see in other forms of animal communication, but limitedly. Let me explain a little bit. There's no question that all animals have some form of communication to express their needs and basic desires; that's not in-argument. The question is: Are they be able to talk about things, not in front of themselves? Are they able to talk about hypotheticals, or make suggestions? Are they able to talk about things that happened in the past, or what may happen in the future?

Maybe. This is why I’m going to say maybe. We know, for example, that most species of birds are able to describe where food or mates are when they're not in front of them—as if to say, “Oh yeah, that flower, that is about five miles that way. That's a really good place to go get nectar.” We know that birds and bees tend to do this, and we see other mammals, especially other primates, have aspects to this in their forms of communication. However, we're still not sure what they actually do among their species.

We are not talking about mimicking human language, and that's a really crucial piece of this argument. We are not talking about when we try to teach a parrot how to talk, or when we try and force another primate to learn a primary sign language. That is not how they communicate with each other, and so we have to abolish that concept entirely.

Because we understand that what a different species used to communicate with its peers is going to be different than what humans do, the research that I’m referring to requires analyzing, observing and being descriptive about what species do amongst their own kind, and to a lesser extent to other animals in the region.

When we talk about animal communication, I love this old The Far Side comic—I'm sorry, I’m a Gen X and The Far Side was part of my upbringing.

Far Side Comic showing what a human is saying to their dog, and that the dog only understands their name.

It encapsulates everything that we think of about animal communication versus human language, as far as what we say to them versus what they hear and understand. I absolutely love and adore The Far Side , especially this comic but I’ll give you an example in real life. I have a cat her name is Bella and she is 16. When I think about my cat, and I’ve had her since she was a kitten since she was three months old, there's a ton of things that we communicate to each other through voice and through body language. She tells me when she needs attention and love, and when she thinks I need attention and love. She tells me very clearly when she has no food in her bowl or its old food and it's not acceptable anymore. She plays well, not so much anymore, but certainly when she was younger, and she definitely communicates that when I go away, she doesn't like that, and when I come home, she makes that very well known. I communicate to her when she does it behavior that I do not approve of, like if she were to scratch the furniture—which she's never done save for once, and it was to get my attention because I forgot to feed her, so she knows how to get my attention. We have a form of communication. When I am at a low point, she's one of those folks I confide in; I cry and she's there. She cuddles me, and I cuddle her; I tell her all my hopes, fears, desires and wishes. and she purrs.

Now, does she understand what I’m saying? Or, is she like The Far Side comic where she just hears noise and she doesn't know what it is? I don't speak cat and she doesn't speak human, so I don't know what really is able to be communicated as far as displacement, as far as productivity or creativity. We certainly have arbitrary sounds and meanings for those sounds. It has often been said, the cats probably learn to meow because of humans their interactions with humans. When they meow in certain ways, humans do certain activities, and it's a symbiotic relationship. I think there's some of that that's true. But she's not able to tell me her deepest hopes, wishes and desires; she's not able to tell me what she thinks might happen in the future, or what did happen in the past. I don't know what she thinks really, although as I’m saying this, she's walking around my feet right now, because she's clearly telling me she doesn't want me talking like this, she doesn't want the camera and she doesn't want the lights. She wants me on the bed right now; she's able to communicate her needs and basic desires. But not much more. Is that to say that she can do that with a different cat? Who's to say?

Where we have been starting to observe a few pieces with respect to animal communication and whether they might have a language, primarily, has to do with our primate cousins. We know certain things to be true. First of all, their vocal tracks are not like human vocal tracks; they are well more primitive, to the point that they cannot produce the sounds that we can produce. We know that part that goes out the window. Yes, it is true that, certainly for other great apes like chimpanzees and gorillas, some have been taught American Sign Language, in particular, and a few other primary sign languages. But—and this is a huge ‘but’—their learning is very slow and formulaic, and they basically get stuck at the level of a three-year-old. If you remember telegraphic speech from child language acquisition in the previous chapter, they're not able to do much more than that, at least not in ASL. They're also not able to create with ASL very well at all. Therefore, I would argue that you can throw out using any kind of human language with a primate; it's probably not going to work.

All that being said, there is quite a bit of evidence to suggest that they might have something primitive. I’m saying the term ‘primitive’, but I do not want you to think that this is a prescriptive use of it. It's saying that this is a very early stage, and maybe in a millennium or several they might have the capability to use a language, much like a human language. At this stage, we don't know. What do we know is that chimpanzees and other great apes are able to teach each other tools. Chimpanzees are particularly good at this, but even we see this in gorillas and some other great apes. We also know that other primates use sounds to communicate things beyond basic needs and desires, not just a warning system, not just to say, “Hey, I need food” or “Hey, I need sex.” You do observe them using the sounds in more arbitrary ways. But—and this is an incredibly important point—we are still trying to decipher what those calls and sounds mean. When we observe our primate cousins teaching each other how to use tools, they are not necessarily using a vocal communication to do it. There is some kind of gesturing. I don't really want to call a sign language yet, because I think it's too early to say that, but our colleagues and primatologist are showing us that our primate cousins, especially the great apes are able to use some kind of communication that's at a higher level than what most other animals do.

Primates are an interesting discussion. What is also interesting, and this is in the video below is Zipf’s Law, and the video is going to go a little more into that. Here's the interesting thing: It could be that dolphins in particular might have a language. You may have heard of studies on dolphin communication before, and this is an area that continuously evolves. Suffice it to say we are very much at the precipice of understanding what other animals do when they need to talk to each other, when they need to communicate to one another, beyond their basic needs, hopes and desires. We are still learning so much about what our own brains do, let alone what other brains of other animals do. So, we'll come back to this—maybe not in this class, and maybe not in the next year, but certainly in the future, so keep an eye on this.

10.1.2 More on Zipf's Law, from NOVA Wonders (PBS)

To finish things off, watch the video below about Zipf's Law, and why we still have so much more to learn about other animals and their methods of communication. (The video is captioned.)

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Animal behavior research is getting better at keeping observer bias from sneaking in – but there’s still room to improve

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Professor and Associate Head of Psychology, University of Tennessee

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Todd M. Freeberg does not work for, consult, own shares in or receive funding from any company or organisation that would benefit from this article, and has disclosed no relevant affiliations beyond their academic appointment.

University of Tennessee provides funding as a member of The Conversation US.

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Animal behavior research relies on careful observation of animals. Researchers might spend months in a jungle habitat watching tropical birds mate and raise their young. They might track the rates of physical contact in cattle herds of different densities. Or they could record the sounds whales make as they migrate through the ocean.

Animal behavior research can provide fundamental insights into the natural processes that affect ecosystems around the globe, as well as into our own human minds and behavior.

I study animal behavior – and also the research reported by scientists in my field. One of the challenges of this kind of science is making sure our own assumptions don’t influence what we think we see in animal subjects. Like all people, how scientists see the world is shaped by biases and expectations, which can affect how data is recorded and reported. For instance, scientists who live in a society with strict gender roles for women and men might interpret things they see animals doing as reflecting those same divisions .

The scientific process corrects for such mistakes over time, but scientists have quicker methods at their disposal to minimize potential observer bias. Animal behavior scientists haven’t always used these methods – but that’s changing. A new study confirms that, over the past decade, studies increasingly adhere to the rigorous best practices that can minimize potential biases in animal behavior research.

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Biases and self-fulfilling prophecies

A German horse named Clever Hans is widely known in the history of animal behavior as a classic example of unconscious bias leading to a false result.

Around the turn of the 20th century , Clever Hans was purported to be able to do math. For example, in response to his owner’s prompt “3 + 5,” Clever Hans would tap his hoof eight times. His owner would then reward him with his favorite vegetables. Initial observers reported that the horse’s abilities were legitimate and that his owner was not being deceptive.

However, careful analysis by a young scientist named Oskar Pfungst revealed that if the horse could not see his owner, he couldn’t answer correctly. So while Clever Hans was not good at math, he was incredibly good at observing his owner’s subtle and unconscious cues that gave the math answers away.

In the 1960s, researchers asked human study participants to code the learning ability of rats. Participants were told their rats had been artificially selected over many generations to be either “bright” or “dull” learners. Over several weeks, the participants ran their rats through eight different learning experiments.

In seven out of the eight experiments , the human participants ranked the “bright” rats as being better learners than the “dull” rats when, in reality, the researchers had randomly picked rats from their breeding colony. Bias led the human participants to see what they thought they should see.

Eliminating bias

Given the clear potential for human biases to skew scientific results, textbooks on animal behavior research methods from the 1980s onward have implored researchers to verify their work using at least one of two commonsense methods.

One is making sure the researcher observing the behavior does not know if the subject comes from one study group or the other. For example, a researcher would measure a cricket’s behavior without knowing if it came from the experimental or control group.

The other best practice is utilizing a second researcher, who has fresh eyes and no knowledge of the data, to observe the behavior and code the data. For example, while analyzing a video file, I count chickadees taking seeds from a feeder 15 times. Later, a second independent observer counts the same number.

Yet these methods to minimize possible biases are often not employed by researchers in animal behavior, perhaps because these best practices take more time and effort.

In 2012, my colleagues and I reviewed nearly 1,000 articles published in five leading animal behavior journals between 1970 and 2010 to see how many reported these methods to minimize potential bias. Less than 10% did so. By contrast, the journal Infancy, which focuses on human infant behavior, was far more rigorous: Over 80% of its articles reported using methods to avoid bias.

It’s a problem not just confined to my field. A 2015 review of published articles in the life sciences found that blind protocols are uncommon . It also found that studies using blind methods detected smaller differences between the key groups being observed compared to studies that didn’t use blind methods, suggesting potential biases led to more notable results.

In the years after we published our article, it was cited regularly and we wondered if there had been any improvement in the field. So, we recently reviewed 40 articles from each of the same five journals for the year 2020.

We found the rate of papers that reported controlling for bias improved in all five journals , from under 10% in our 2012 article to just over 50% in our new review. These rates of reporting still lag behind the journal Infancy, however, which was 95% in 2020.

All in all, things are looking up, but the animal behavior field can still do better. Practically, with increasingly more portable and affordable audio and video recording technology, it’s getting easier to carry out methods that minimize potential biases. The more the field of animal behavior sticks with these best practices, the stronger the foundation of knowledge and public trust in this science will become.

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Disclaimer: Early release articles are not considered as final versions. Any changes will be reflected in the online version in the month the article is officially released.

Volume 30, Number 7—July 2024

Highly Pathogenic Avian Influenza A(H5N1) Clade Virus Infection in Domestic Dairy Cattle and Cats, United States, 2024

Suggested citation for this article

We report highly pathogenic avian influenza A(H5N1) virus in dairy cattle and cats in Kansas and Texas, United States, which reflects the continued spread of clade viruses that entered the country in late 2021. Infected cattle experienced nonspecific illness, reduced feed intake and rumination, and an abrupt drop in milk production, but fatal systemic influenza infection developed in domestic cats fed raw (unpasteurized) colostrum and milk from affected cows. Cow-to-cow transmission appears to have occurred because infections were observed in cattle on Michigan, Idaho, and Ohio farms where avian influenza virus–infected cows were transported. Although the US Food and Drug Administration has indicated the commercial milk supply remains safe, the detection of influenza virus in unpasteurized bovine milk is a concern because of potential cross-species transmission. Continued surveillance of highly pathogenic avian influenza viruses in domestic production animals is needed to prevent cross-species and mammal-to-mammal transmission.

Highly pathogenic avian influenza (HPAI) viruses pose a threat to wild birds and poultry globally, and HPAI H5N1 viruses are of even greater concern because of their frequent spillover into mammals. In late 2021, the Eurasian strain of H5N1 (clade was detected in North America ( 1 , 2 ) and initiated an outbreak that continued into 2024. Spillover detections and deaths from this clade have been reported in both terrestrial and marine mammals in the United States ( 3 , 4 ). The detection of HPAI H5N1 clade virus in severe cases of human disease in Ecuador ( 5 ) and Chile ( 6 ) raises further concerns regarding the pandemic potential of specific HPAI viruses.

In February 2024, veterinarians were alerted to a syndrome occurring in lactating dairy cattle in the panhandle region of northern Texas. Nonspecific illness accompanied by reduced feed intake and rumination and an abrupt drop in milk production developed in affected animals. The milk from most affected cows had a thickened, creamy yellow appearance similar to colostrum. On affected farms, incidence appeared to peak 4–6 days after the first animals were affected and then tapered off within 10–14 days; afterward, most animals were slowly returned to regular milking. Clinical signs were commonly reported in multiparous cows during middle to late lactation; ≈10%–15% illness and minimal death of cattle were observed on affected farms. Initial submissions of blood, urine, feces, milk, and nasal swab samples and postmortem tissues to regional diagnostic laboratories did not reveal a consistent, specific cause for reduced milk production. Milk cultures were often negative, and serum chemistry testing showed mildly increased aspartate aminotransferase, gamma-glutamyl transferase, creatinine kinase, and bilirubin values, whereas complete blood counts showed variable anemia and leukocytopenia.

In early March 2024, similar clinical cases were reported in dairy cattle in southwestern Kansas and northeastern New Mexico; deaths of wild birds and domestic cats were also observed within affected sites in the Texas panhandle. In > 1 dairy farms in Texas, deaths occurred in domestic cats fed raw colostrum and milk from sick cows that were in the hospital parlor. Antemortem clinical signs in affected cats were depressed mental state, stiff body movements, ataxia, blindness, circling, and copious oculonasal discharge. Neurologic exams of affected cats revealed the absence of menace reflexes and pupillary light responses with a weak blink response.

On March 21, 2024, milk, serum, and fresh and fixed tissue samples from cattle located in affected dairies in Texas and 2 deceased cats from an affected Texas dairy farm were received at the Iowa State University Veterinary Diagnostic Laboratory (ISUVDL; Ames, IA, USA). The next day, similar sets of samples were received from cattle located in affected dairies in Kansas. Milk and tissue samples from cattle and tissue samples from the cats tested positive for influenza A virus (IAV) by screening PCR, which was confirmed and characterized as HPAI H5N1 virus by the US Department of Agriculture National Veterinary Services Laboratory. Detection led to an initial press release by the US Department of Agriculture Animal and Plant Health Inspection Service on March 25, 2024, confirming HPAI virus in dairy cattle ( 7 ). We report the characterizations performed at the ISUVDL for HPAI H5N1 viruses infecting cattle and cats in Kansas and Texas.

Materials and Methods

Milk samples (cases 2–5) and fresh and formalin-fixed tissues (cases 1, 3–5) from dairy cattle were received at the ISUVDL from Texas on March 21 and from Kansas on March 22, 2024. The cattle exhibited nonspecific illness and reduced lactation, as described previously. The tissue samples for diagnostic testing came from 3 cows that were euthanized and 3 that died naturally; all postmortem examinations were performed on the premises of affected farms.

The bodies of 2 adult domestic shorthaired cats from a north Texas dairy farm were received at the ISUVDL for a complete postmortem examination on March 21, 2024. The cats were found dead with no apparent signs of injury and were from a resident population of ≈24 domestic cats that had been fed milk from sick cows. Clinical disease in cows on that farm was first noted on March 16; the cats became sick on March 17, and several cats died in a cluster during March 19–20. In total, >50% of the cats at that dairy became ill and died. We collected cerebrum, cerebellum, eye, lung, heart, spleen, liver, lymph node, and kidney tissue samples from the cats and placed them in 10% neutral-buffered formalin for histopathology.

At ISUVDL, we trimmed, embedded in paraffin, and processed formalin-fixed tissues from affected cattle and cats for hematoxylin/eosin staining and histologic evaluation. For immunohistochemistry (IHC), we prepared 4-µm–thick sections from paraffin-embedded tissues, placed them on Superfrost Plus slides (VWR, https://www.vwr.com ), and dried them for 20 minutes at 60°C. We used a Ventana Discovery Ultra IHC/ISH research platform (Roche, https://www.roche.com ) for deparaffinization until and including counterstaining. We obtained all products except the primary antibody from Roche. Automated deparaffination was followed by enzymatic digestion with protease 1 for 8 minutes at 37°C and endogenous peroxidase blocking. We obtained the primary influenza A virus antibody from the hybridoma cell line H16-L10–4R5 (ATCC, https://www.atcc.org ) and diluted at 1:100 in Discovery PSS diluent; we incubated sections with antibody for 32 minutes at room temperature. Next, we incubated the sections with a hapten-labeled conjugate, Discovery anti-mouse HQ, for 16 minutes at 37°C followed by a 16-minute incubation with the horse radish peroxidase conjugate, Discovery anti-HQ HRP. We used a ChromoMap DAB kit for antigen visualization, followed by counterstaining with hematoxylin and then bluing. Positive controls were sections of IAV-positive swine lung. Negative controls were sections of brain, lung, and eyes from cats not infected with IAV.

We diluted milk samples 1:3 vol/vol in phosphate buffered saline, pH 7.4 (Gibco/Thermo Fisher Scientific, https://www.thermofisher.com ) by mixing 1 unit volume of milk and 3 unit volumes of phosphate buffered saline. We prepared 10% homogenates of mammary glands, brains, lungs, spleens, and lymph nodes in Earle’s balanced salt solution (Sigma-Aldrich, https://www.sigmaaldrich.com ). Processing was not necessary for ocular fluid, rumen content, or serum samples. After processing, we extracted samples according to a National Animal Health Laboratory Network (NAHLN) protocol that had 2 NAHLN-approved deviations for ISUVDL consisting of the MagMax Viral RNA Isolation Kit for 100 µL sample volumes and a Kingfisher Flex instrument (both Thermo Fisher Scientific).

We performed real-time reverse transcription PCR (rRT-PCR) by using an NAHLN-approved assay with 1 deviation, which was the VetMAX-Gold SIV Detection kit (Thermo Fisher Scientific), to screen for the presence of IAV RNA. We tested samples along with the VetMAX XENO Internal Positive Control to monitor the possible presence of PCR inhibitors. Each rRT-PCR 96-well plate had 2 positive amplification controls, 2 negative amplification controls, 1 positive extraction control, and 1 negative extraction control. We ran the rRT-PCR on an ABI 7500 Fast thermocycler and analyzed data with Design and Analysis Software 2.7.0 (both Thermo Fisher Scientific). We considered samples with cycle threshold (Ct) values <40.0 to be positive for virus.

After the screening rRT-PCR, we analyzed IAV RNA–positive samples for the H5 subtype and H5 clade by using the same RNA extraction and NAHLN-approved rRT-PCR protocols as described previously, according to standard operating procedures. We performed PCR on the ABI 7500 Fast thermocycler by using appropriate controls to detect H5-specific IAV. We considered samples with Ct values <40.0 to be positive for the IAV H5 subtype.

We conducted genomic sequencing of 2 milk samples from infected dairy cattle from Texas and 2 tissue samples (lung and brain) from cats that died at a different Texas dairy. We subjected the whole-genome sequencing data to bioinformatics analysis to assemble the 8 different IAV segment sequences according to previously described methods ( 8 ). We used the hemagglutinin (HA) and neuraminidase (NA) sequences for phylogenetic analysis. We obtained reference sequences for the HA and NA segments of IAV H5 clade from publicly available databases, including GISAID ( https://www.gisaid.org ) and GenBank. We aligned the sequences by using MAFFT version 7.520 software ( https://mafft.cbrc.jp/alignment/server/index.html ) to create multiple sequence alignments for subsequent phylogenetic analysis. We used IQTree2 ( https://github.com/iqtree/iqtree2 ) to construct the phylogenetic tree from the aligned sequences. The software was configured to automatically identify the optimal substitution model by using the ModelFinder Plus option, ensuring the selection of the most suitable model for the dataset and, thereby, improving the accuracy of the reconstructed tree. We visualized the resulting phylogenetic tree by using iTOL ( https://itol.embl.de ), a web-based platform for interactive tree exploration and annotation.

Gross Lesions in Cows and Cats

All cows were in good body condition with adequate rumen fill and no external indications of disease. Postmortem examinations of the affected dairy cows revealed firm mammary glands typical of mastitis; however, mammary gland lesions were not consistent. Two cows that were acutely ill before postmortem examination had grossly normal milk and no abnormal mammary gland lesions. The gastrointestinal tract of some cows had small abomasal ulcers and shallow linear erosions of the intestines, but those observations were also not consistent in all animals. The colon contents were brown and sticky, suggesting moderate dehydration. The feces contained feed particles that appeared to have undergone minimal ruminal fermentation. The rumen contents had normal color and appearance but appeared to have undergone minimal fermentation.

The 2 adult cats (1 intact male, 1 intact female) received at the ISUVDL were in adequate body and postmortem condition. External examination was unremarkable. Mild hemorrhages were observed in the subcutaneous tissues over the dorsal skull, and multifocal meningeal hemorrhages were observed in the cerebrums of both cats. The gastrointestinal tracts were empty, and no other gross lesions were observed.

Microscopic Lesions in Cows and Cats

Mammary gland lesions in cattle in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. A, B) Mammary gland tissue sections stained with hematoxylin and eosin. A) Arrowheads indicate segmental loss within open secretory mammary alveoli. Original magnification ×40. B) Arrowheads indicate epithelial degeneration and necrosis lining alveoli with intraluminal sloughing. Asterisk indicates intraluminal neutrophilic inflammation. Original magnification ×400. C, D) Mammary gland tissue sections stained by using avian influenza A immunohistochemistry. C) Brown staining indicates lobular distribution of avian influenza A virus. Original magnification ×40. D) Brown staining indicates strong nuclear and intracytoplasmic immunoreactivity of intact and sloughed epithelial cells within mammary alveoli. Original magnification ×400.

Figure 1 . Mammary gland lesions in cattle in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. A, B) Mammary gland...

The chief microscopic lesion observed in affected cows was moderate acute multifocal neutrophilic mastitis ( Figure 1 ); however, mammary glands were not received from every cow. Three cows had mild neutrophilic or lymphocytic hepatitis. Because they were adult cattle, other observed microscopic lesions (e.g., mild lymphoplasmacytic interstitial nephritis and mild to moderate lymphocytic abomasitis) were presumed to be nonspecific, age-related changes. We did not observe major lesions in the other evaluated tissues. We performed IHC for IAV antigen on all evaluated tissues; the only tissues with positive immunoreactivity were mastitic mammary glands from 2 cows that showed nuclear and cytoplasmic labeling of alveolar epithelial cells and cells within lumina ( Figure 1 ) and multifocal germinal centers within a lymph node from 1 cow ( Table 1 ).

Lesions in cat tissues in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Tissue sections were stained with hematoxylin and eosin; insets show brown staining of avian influenza A viruses via immunohistochemistry by using the chromogen 3,3′-diaminobenzidine tetrahydrochloride. Original magnification ×200 for all images and insets. A) Section from cerebral tissue. Arrowheads show perivascular lymphocytic encephalitis, gliosis, and neuronal necrosis. Inset shows neurons. B) Section of lung tissue showing lymphocytic and fibrinous interstitial pneumonia with septal necrosis and alveolar edema; arrowheads indicate lymphocytes. Inset shows bronchiolar epithelium, necrotic cells, and intraseptal mononuclear cells. C) Section of heart tissue. Arrowhead shows interstitial lymphocytic myocarditis and focal peracute myocardial coagulative necrosis. Inset shows cardiomyocytes. D) Section of retinal tissue. Arrowheads show perivascular lymphocytic retinitis with segmental neuronal loss and rarefaction in the ganglion cell layer. Asterisks indicate attenuation of the inner plexiform and nuclear layers with artifactual retinal detachment. Insets shows all layers of the retina segmentally within affected areas have strong cytoplasmic and nuclear immunoreactivity to influenza A virus.

Figure 2 . Lesions in cat tissues in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Tissue sections were stained with...

Both cats had microscopic lesions consistent with severe systemic virus infection, including severe subacute multifocal necrotizing and lymphocytic meningoencephalitis with vasculitis and neuronal necrosis, moderate subacute multifocal necrotizing and lymphocytic interstitial pneumonia, moderate to severe subacute multifocal necrotizing and lymphohistiocytic myocarditis, and moderate subacute multifocal lymphoplasmacytic chorioretinitis with ganglion cell necrosis and attenuation of the internal plexiform and nuclear layers ( Table 2 ; Figure 2 ). We performed IHC for IAV antigen on multiple tissues (brain, eye, lung, heart, spleen, liver, and kidney). We detected positive IAV immunoreactivity in brain (intracytoplasmic, intranuclear, and axonal immunolabeling of neurons), lung, and heart, and multifocal and segmental immunoreactivity within all layers of the retina ( Figure 2 ).

PCR Data from Cows and Cats

We tested various samples from 8 clinically affected mature dairy cows by IAV screening and H5 subtype-specific PCR ( Table 3 ). Milk and mammary gland homogenates consistently showed low Ct values: 12.3–16.9 by IAV screening PCR, 17.6–23.1 by H5 subtype PCR, and 14.7–20.0 by H5 clade PCR (case 1, cow 1; case 2, cows 1 and 2; case 3, cow 1; and case 4, cow 1). We forwarded the samples to the National Veterinary Services Laboratory, which confirmed the virus was an HPAI H5N1 virus strain.

When available, we also tested tissue homogenates (e.g., lung, spleen, and lymph nodes), ocular fluid, and rumen contents from 6 cows by IAV and H5 subtype-specific PCR ( Table 3 ). However, the PCR findings were not consistent. For example, the tissue homogenates and ocular fluid tested positive in some but not all cows. In case 5, cow 1, the milk sample tested negative by IAV screening PCR, but the spleen homogenate tested positive by IAV screening, H5 subtype, and H5 PCR. For 2 cows (case 3, cow 1; and case 4, cow 1) that had both milk and rumen contents available, both samples tested positive for IAV. Nevertheless, all IAV-positive nonmammary gland tissue homogenates, ocular fluid, and rumen contents had markedly elevated Ct values in contrast to the low Ct values for milk and mammary gland homogenate samples.

We tested brain and lung samples from the 2 cats (case 6, cats 1 and 2) by IAV screening and H5 subtype-specific PCR ( Table 3 ). Both sample types were positive by IAV screening PCR; Ct values were 9.9–13.5 for brain and 17.4–24.4 for lung samples, indicating high amounts of virus nucleic acid in those samples. The H5 subtype and H5 PCR results were also positive for the brain and lung samples; Ct values were consistent with the IAV screening PCR ( Table 3 ).

Phylogenetic Analyses

We assembled the sequences of all 8 segments of the HPAI viruses from both cow milk and cat tissue samples. We used the hemagglutinin (HA) and neuraminidase (NA) sequences specifically for phylogenetic analysis to delineate the clade of the HA gene and subtype of the NA gene.

Phylogenetic analysis of hemagglutinin gene sequences in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Colors indicate different clades. Red text indicates the virus gene sequences from bovine milk and cats described in this report, confirming those viruses are highly similar and belong to H5 clade The hemagglutinin sequences from this report are most closely related to A/avian/Guanajuato/CENAPA-18539/2023|EPI_ISL_18755544|A_/_H5 (GISAID, https://www.gisaid.org) and have 99.66%–99.72% nucleotide identities.

Figure 3 . Phylogenetic analysis of hemagglutinin gene sequences in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Colors indicate different...

For HA gene analysis, both HA sequences derived from cow milk samples exhibited a high degree of similarity, sharing 99.88% nucleotide identity, whereas the 2 HA sequences from cat tissue samples showed complete identity at 100%. The HA sequences from the milk samples had 99.94% nucleotide identities with HA sequences from the cat tissues, resulting in a distinct subcluster comprising all 4 HA sequences, which clustered together with other H5N1 viruses belonging to clade ( Figure 3 ). The HA sequences were deposited in GenBank (accession nos. PP599465 [case 2, cow 1], PP599473 [case 2, cow 2], PP692142 [case 6, cat 1], and PP692195 [case 6, cat 2]).

Phylogenetic analysis of neuraminidase gene sequences in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Colors indicate different subtypes. Red text indicates the virus gene sequences from bovine milk and cats described in this report, confirming those viruses belong to the N1 subtype. The neuraminidase sequences from this report had 99.52%–99.59% nucleotide identities to sequences from viruses isolated from a chicken and wild birds in 2023.

Figure 4 . Phylogenetic analysis of neuraminidase gene sequences in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Colors indicate different...

For NA gene analysis, the 2 NA sequences obtained from cow milk samples showed 99.93% nucleotide identity. Moreover, the NA sequences derived from the milk samples exhibited complete nucleotide identities (100%) with those from the cat tissues. The 4 NA sequences were grouped within the N1 subtype of HPAI viruses ( Figure 4 ). The NA sequences were deposited in GenBank (accession nos. PP599467 [case 2, cow 1], PP599475 [case 2, cow 2], PP692144 [case 6, cat 1], and PP692197 [case 6, cat 2]).

This case series differs from most previous reports of IAV infection in bovids, which indicated cattle were inapparently infected or resistant to infection ( 9 ). We describe an H5N1 strain of IAV in dairy cattle that resulted in apparent systemic illness, reduced milk production, and abundant virus shedding in milk. The magnitude of this finding is further emphasized by the high death rate (≈50%) of cats on farm premises that were fed raw colostrum and milk from affected cows; clinical disease and lesions developed that were consistent with previous reports of H5N1 infection in cats presumably derived from consuming infected wild birds ( 10 – 12 ). Although exposure to and consumption of dead wild birds cannot be completely ruled out for the cats described in this report, the known consumption of unpasteurized milk and colostrum from infected cows and the high amount of virus nucleic acid within the milk make milk and colostrum consumption a likely route of exposure. Therefore, our findings suggest cross-species mammal-to-mammal transmission of HPAI H5N1 virus and raise new concerns regarding the potential for virus spread within mammal populations. Horizontal transmission of HPAI H5N1 virus has been previously demonstrated in experimentally infected cats ( 13 ) and ferrets ( 14 ) and is suspected to account for large dieoffs observed during natural outbreaks in mink ( 15 ) and sea lions ( 16 ). Future experimental studies of HPAI H5N1 virus in dairy cattle should seek to confirm cross-species transmission to cats and potentially other mammals.

Clinical IAV infection in cattle has been infrequently reported in the published literature. The first report occurred in Japan in 1949, where a short course of disease with pyrexia, anorexia, nasal discharge, pneumonia, and decreased lactation developed in cattle ( 17 ). In 1997, a similar condition occurred in dairy cows in southwest England leading to a sporadic drop in milk production ( 18 ), and IAV seroconversion was later associated with reduced milk yield and respiratory disease ( 19 – 21 ). Rising antibody titers against human-origin influenza A viruses (H1N1 and H3N2) were later again reported in dairy cattle in England, which led to an acute fall in milk production during October 2005–March 2006 ( 22 ). Limited reports of IAV isolation from cattle exist; most reports occurred during the 1960s and 1970s in Hungary and in the former Soviet Union, where H3N2 was recovered from cattle experiencing respiratory disease ( 9 , 23 ). Direct detection of IAV in milk and the potential transmission from cattle to cats through feeding of unpasteurized milk has not been previously reported.

An IAV-associated drop in milk production in dairy cattle appears to have occurred during > 4 distinct periods and within 3 widely separated geographic areas: 1949 in Japan ( 17 ), 1997–1998 and 2005–2006 in Europe ( 19 , 21 ), and 2024 in the United States (this report). The sporadic occurrence of clinical disease in dairy cattle worldwide might be the result of changes in subclinical infection rates and the presence or absence of sufficient baseline IAV antibodies in cattle to prevent infection. Milk IgG, lactoferrin, and conglutinin have also been suggested as host factors that might reduce susceptibility of bovids to IAV infection ( 9 ). Contemporary estimates of the seroprevalence of IAV antibodies in US cattle are not well described in the published literature. One retrospective serologic survey in the United States in the late 1990s showed 27% of serum samples had positive antibody titers and 31% had low-positive titers for IAV H1 subtype-specific antigen in cattle with no evidence of clinical infections ( 24 ). Antibody titers for H5 subtype-specific antigen have not been reported in US cattle.

The susceptibility of domestic cats to HPAI H5N1 is well-documented globally ( 10 – 12 , 25 – 28 ), and infection often results in neurologic signs in affected felids and other terrestrial mammals ( 4 ). Most cases in cats result from consuming infected wild birds or contaminated poultry products ( 12 , 27 ). The incubation period in cats is short; clinical disease is often observed 2–3 days after infection ( 28 ). Brain tissue has been suggested as the best diagnostic sample to confirm HPAI virus infection in cats ( 10 ), and our results support that finding. One unique finding in the cats from this report is the presence of blindness and microscopic lesions of chorioretinitis. Those results suggest that further investigation into potential ocular manifestations of HPAI H5N1 virus infection in cats might be warranted.

The genomic sequencing and subsequent analysis of clinical samples from both bovine and feline sources provided considerable insights. The HA and NA sequences derived from both bovine milk and cat tissue samples from different Texas farms had a notable degree of similarity. Those findings strongly suggest a shared origin for the viruses detected in the dairy cattle and cat tissues. Further research, case series investigations, and surveillance data are needed to better understand and inform measures to curtail the clinical effects, shedding, and spread of HPAI viruses among mammals. Although pasteurization of commercial milk mitigates risks for transmission to humans, a 2019 US consumer study showed that 4.4% of adults consumed raw milk > 1 time during the previous year ( 29 ), indicating a need for public awareness of the potential presence of HPAI H5N1 viruses in raw milk.

Ingestion of feed contaminated with feces from wild birds infected with HPAI virus is presumed to be the most likely initial source of infection in the dairy farms. Although the exact source of the virus is unknown, migratory birds (Anseriformes and Charadriiformes) are likely sources because the Texas panhandle region lies in the Central Flyway, and those birds are the main natural reservoir for avian influenza viruses ( 30 ). HPAI H5N1 viruses are well adapted to domestic ducks and geese, and ducks appear to be a major reservoir ( 31 ); however, terns have also emerged as an important source of virus spread ( 32 ). The mode of transmission among infected cattle is also unknown; however, horizontal transmission has been suggested because disease developed in resident cattle herds in Michigan, Idaho, and Ohio farms that received infected cattle from the affected regions, and those cattle tested positive for HPAI H5N1 ( 33 ). Experimental studies are needed to decipher the transmission routes and pathogenesis (e.g., replication sites and movement) of the virus within infected cattle.

In conclusion, we showed that dairy cattle are susceptible to infection with HPAI H5N1 virus and can shed virus in milk and, therefore, might potentially transmit infection to other mammals via unpasteurized milk. A reduction in milk production and vague systemic illness were the most commonly reported clinical signs in affected cows, but neurologic signs and death rapidly developed in affected domestic cats. HPAI virus infection should be considered in dairy cattle when an unexpected and unexplained abrupt drop in feed intake and milk production occurs and for cats when rapid onset of neurologic signs and blindness develop. The recurring nature of global HPAI H5N1 virus outbreaks and detection of spillover events in a broad host range is concerning and suggests increasing virus adaptation in mammals. Surveillance of HPAI viruses in domestic production animals, including cattle, is needed to elucidate influenza virus evolution and ecology and prevent cross-species transmission.

Dr. Burrough is a professor and diagnostic pathologist at the Iowa State University College of Veterinary Medicine and Veterinary Diagnostic Laboratory. His research focuses on infectious diseases of livestock with an emphasis on swine.


We thank the faculty and staff at the ISUVDL who contributed to the processing and analysis of clinical samples in this investigation, the veterinarians involved with clinical assessments at affected dairies and various conference calls in the days before diagnostic submissions that ultimately led to the detection of HPAI virus in the cattle, and the US Department of Agriculture National Veterinary Services Laboratory and NAHLN for their roles and assistance in providing their expertise, confirmatory diagnostic support, and communications surrounding the HPAI virus cases impacting lactating dairy cattle.

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  • Figure 1 . Mammary gland lesions in cattle in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. A, B) Mammary...
  • Figure 2 . Lesions in cat tissues in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Tissue sections were stained...
  • Figure 3 . Phylogenetic analysis of hemagglutinin gene sequences in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Colors indicate...
  • Figure 4 . Phylogenetic analysis of neuraminidase gene sequences in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Colors indicate...
  • Table 1 . Microscopic lesions observed in cattle in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024
  • Table 2 . Microscopic lesions observed in cats in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024
  • Table 3 . PCR results from various specimens in study of highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024

Suggested citation for this article : Burrough ER, Magstadt DR, Petersen B, Timmermans SJ, Gauger PC, Zhang J, et al. Highly pathogenic avian influenza A(H5N1) clade virus infection in domestic dairy cattle and cats, United States, 2024. Emerg Infect Dis. 2024 Jul [ date cited ]. https://doi.org/10.3201/eid3007.240508

DOI: 10.3201/eid3007.240508

Original Publication Date: April 29, 2024

Table of Contents – Volume 30, Number 7—July 2024

Please use the form below to submit correspondence to the authors or contact them at the following address:

Eric R. Burrough, Iowa State University Veterinary Diagnostic Laboratory, 1937 Christensen Dr, Ames, IA 50011, USA

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    Our community of horse doctors connects you to more than 9,000 veterinarians and veterinary students who make a difference every day in horse health, just like you!

  29. Deep sea mining could be disastrous for marine animals

    In a recent study published in Deep-Sea Research Part I: Oceanographic Research Papers, researchers of Wageningen University & Research and the University of Bergen have shown that release of deep ...

  30. Volume 30, Number 7—July 2024

    Research Highly Pathogenic Avian Influenza A(H5N1) Clade Virus Infection in Domestic Dairy Cattle and Cats, United States, 2024 ... for ocular fluid, rumen content, or serum samples. After processing, we extracted samples according to a National Animal Health Laboratory Network (NAHLN) protocol that had 2 NAHLN-approved deviations for ...