Johan Lind

• Senior Associate Professor at Linköping University

In speech, sequences of speech sounds (or phonemes) are combined into words and sentences. In recent literature, great apes have been shown to possess several vocal capacities previously believed to elude them, including voluntary control of the articulators, and a capacity to produce several speech-like sounds. We identify a core constraint on syl...

Developmental psychology and cultural evolution are concerned with the same research questions but rarely interact. Collaboration between these fields could lead to substantial progress. Developmental psychology and related fields such as educational science and linguistics explore how behavior and cognition develop through combinations of social a...

Human language is unique in its compositional, open-ended, and sequential form, and its evolution is often solely explained by advantages of communication. However, it has proven challenging to identify an evolutionary trajectory from a world without language to a world with language, especially while at the same time explaining why such an advanta...

This chapter explains how the world offers many resources to organisms, noting that obtaining resources is hard. It covers exploration that takes time, behavior that may have delayed consequences, and informative stimuli that are often mixed with noninformative ones. It also mentions animals that have evolved several solutions to challenges, such a...

This chapter shows that cultural evolution enables the accumulation of vast amounts of knowledge about the world, much beyond what would be possible for individuals. It emphasizes that the availability of cultural information has momentous consequences for the behavioral and cognitive architecture of individuals and that the presence of culture all...

This chapter attempts to draw general conclusions about animal intelligence. It builds on certain hypotheses about animal intelligence to suggest a standard model that recognizes the power of associative learning, the costs of behavioral flexibility, and the need for genetic guidance of learning and decision making. It also explains empirical obser...

This chapter presents seven hypotheses about human and animal intelligence. It talks about recent advances in machine learning that have led to a new understanding of animal intelligence in terms of associative learning, although sometimes its at odds with common views. It also investigates animal intelligence that consists of a sophisticated inter...

This chapter focuses on the necessity of genetic guidance for productive learning and explores how genetic predispositions can influence choice of behavior, exploratory tendencies, and what an animal learns about. It describes how genetic predispositions foster efficient behavioral adaptations, and limit what can be learned and can be maladaptive o...

This book offers a unified view of the evolution of intelligence, presenting a bold and provocative new account of how animals and humans have followed two powerful yet very different evolutionary paths to intelligence. The book shows how animals rely on robust associative mechanisms that are guided by genetic information, which enable animals to s...

This chapter discusses in some detail the costs and benefits of various ways of transmitting sequential information. It includes social learning of behavioral sequences from observation, learning about sequential episodic and semantic information, transmission of sequential information through language, and teaching. It also points out how sequenti...

This chapter explores two important consequences of the genetic abilities that started the human evolutionary transition: faithful sequence representation and mental flexibility. The chapter emphasizes how genetic abilities have profound consequences for the behavioral and mental skills of humans and for the cultural evolutionary process. It also s...

This chapter points out how decision making, whether in animals, humans, or machines, combines information about the present situation with information stored in memory. It surveys what information animals memorize and use, recalling that animals are often very skilled in using information from sense organs and internal stimuli. It also explores ho...

This chapter argues that, during human evolution, a new, eventually more effective strategy has emerged to tackle combinatorial dilemmas. The chapter explores the evolutionary transition of relying on cultural knowledge rather than genetic knowledge, which has fundamentally changed human evolution. It also makes two claims about the human mental ar...

This chapter explores a major hypothesis about human evolution: uniquely human mental skills are learned, while the way animals learn and decide is essentially hardwired. It surveys the characteristics of human mental skills and leverage findings from developmental psychology in order to argue that the transmission of mental skills is ubiquitous in...

This chapter focuses on aspects of social learning that are deemed relevant to understanding the differences between humans and animals. It summarizes what animals can learn socially and discusses the nature of social information, showing that many observations of social learning can result from associative learning. It also analyzes the limits of...

This chapter discusses a simple yet powerful mental mechanism that may account for how animals learn most behavioral sequences. It refers to this mental mechanism as chaining, which is a term introduced by B.F. Skinner to describe a type of learning in which known behaviors or shorter sequences are linked to form new or longer sequences. The chapte...

This chapter explores how human cooperation is influenced by uniquely human mental skills and how it can increase the ability to discover and execute productive behavioral sequences. It argues that human-level cooperation requires a sophisticated understanding of social relations or a mental map of the individual's social world. It also discusses d...

This chapter considers the evidence of animal learning for more sophisticated mental mechanisms, particularly mechanisms that would qualify as thinking, insight, or mental problem solving. It defines thinking as a recombination of causal information gathered from different experiences that echoes common uses of insight, mental problem solving, and...

This chapter summarizes both behavioral and mental sequential abilities of humans, discussing when these abilities may have appeared and how they develop during an individual's lifetime. The chapter looks at the main hypotheses about human cognitive evolution. It describes uniquely human behavior that includes extensive collaborative sequences, lan...

This chapter argues that behavioral and mental mechanisms in animals are genetically controlled to a much greater extent than in humans. It points out that a human, while remaining human, can learn to be a singer, a surgeon, and countless other things, with each behavioral skill underpinned by a large number of unique, learned mental skills. It als...

Identifying cognitive capacities underlying the human evolutionary transition is challenging, and many hypotheses exist for what makes humans capable of, for example, producing and understanding language, preparing meals, and having culture on a grand scale. Instead of describing processes whereby information is processed, recent studies have sugge...

In a recent study, Wasserman, Kain, and O'Donoghue (Current Biology, 33(6), 1112-1116, 2023) set out to resolve the associative learning paradox by showing that pigeons can solve a complex category learning task through associative learning. The present Outlook paper presents their findings, expands on this paradox, and discusses implications of th...

Abnormal behaviours are common in captive animals, and despite a lot of research, the development, maintenance and alleviation of these behaviours are not fully understood. Here, we suggest that conditioned reinforcement can induce sequential dependencies in behaviour that are difficult to infer from direct observation. We develop this hypothesis u...

Evolution of Learning and Memory Mechanisms is an exploration of laboratory and field research on the many ways that evolution has influenced learning and memory processes, such as associative learning, social learning, and spatial, working, and episodic memory systems. This volume features research by both outstanding early-career scientists as we...

Human language is unique in its compositional and sequential form. Language evolution is often explained by advantages of communication. However, it has proven challenging to identify an evolutionary trajectory from a world without language to a world with language, especially while at the same time explaining why such an advantageous trait has not...

The general process- and adaptive specialization hypotheses represent two contrasting explanations for understanding intelligence in non-human animals. The general process hypothesis proposes that associative learning underlies all learning, whereas the adaptive specialization hypothesis suggests additional distinct learning processes required for...

A widespread and popular belief posits that humans possess a cognitive capacity that is limited to keeping track of and maintaining stable relationships with approximately 150 people. This influential number, 'Dunbar's number', originates from an extrapolation of a regression line describing the relationship between relative neocortex size and grou...

We present a new mathematical formulation of associative learning focused on non-human animals, which we call A-learning. Building on current animal learning theory and machine learning, A-learning is composed of two learning equations, one for stimulus-response values and one for stimulus values (conditioned reinforcement). A third equation implem...

Primate brains differ in size and architecture. Hypotheses to explain this variation are numerous and many tests have been carried out. However, after body size has been accounted for there is little left to explain. The proposed explanatory variables for the residual variation are many and covary, both with each other and with body size. Further,...

Social transmission of information is a key phenomenon in the evolution of behaviour and in the establishment of traditions and culture. The diversity of social learning phenomena has engendered a diverse terminology and numerous ideas about underlying learning mechanisms, at the same time that some researchers have called for a unitary analysis of...

Supplementary materials for “Social learning through associative processes: A computational theory”

There is a new associative learning paradox. The power of associative learning for producing flexible behaviour in non-human animals is downplayed or ignored by researchers in animal cognition, whereas artificial intelligence research shows that associative learning models can beat humans in chess. One phenomenon in which associative learning often...

Primate brains differ in size and architecture. Hypotheses to explain this variation are numerous and many tests have been carried out. However, after body size has been accounted for there is little left to explain. The proposed explanatory variables for the residual variation are many and covary, both with each other and with body size. Further,...

Social transmission of information is a key phenomenon in the evolution of behavior and in the establishment of traditions and culture. The diversity of social learning phenomena has engendered a diverse terminology and numerous ideas about underlying learning mechanisms, at the same time that some researchers have called for a unitary analysis of...

Humans stand out among animals for their unique capacities in domains such as language, culture and imitation, yet it has been difficult to identify cognitive elements that are specifically human. Most research has focused on how information is processed after it is acquired, e.g. in problem solving or ‘insight’ tasks, but we may also look for spec...

Many questions in animal intelligence and cognition research are challenging. One challenge is to identify mechanisms underlying reasoning in experiments. Here, we provide a way to design such tests in non-human animals. We know from research in skill acquisition in humans that reasoning and thinking can take time because some problems are processe...

Experiments inspired by Aesop's fable The crow and the pitcher have been suggested to show that some birds (rooks, Corvus frugilegus, New Caledonian crows, Corvus moneduloides, and Eurasian jays, Garrulus glandarius) understand cause–effect relationships pertaining to water displacement. For example, the birds may prefer to drop stones in water rat...

Behaving efficiently (optimally or near-optimally) is central to animals’ adaptation to their environment. Much evolutionary biology assumes, implicitly or explicitly, that optimal behavioural strategies are genetically inherited, yet the behaviour of many animals depends crucially on learning. The question of how learning contributes to optimal be...

This package contains simulation code accompanying the paper "The power of associative learning and the ontogeny of optimal behavior," by Magnus Enquist, Johan Lind, and Stefano Ghirlanda. The package contains code to reproduce all simulations in the paper, and documentation on how the code works.

We performed a meta-analysis of over 90 data sets from delayed matching-to-sample (DMTS) studies with 25 species (birds, mammals, and bees). In DMTS, a sample stimulus is first presented and then removed. After a delay, two (or more) comparison stimuli are presented, and the subject is rewarded for choosing the one matching the sample. We used data...

Recent studies have shown that large fuel loads in small birds impair ¯ying ability. This is the ®rst study to show how migratory fuel load a€ects ¯ying ability, such as velocity and height gained at take-o€ in a predator escape situation, in a medium-distance migrant , and whether they adjust their take-o€ according to predator attack angle. First...

Across many taxa, intriguing positive correlations exist between intelligence (measured by proxy as encephalization), behavioral repertoire size, and lifespan. Here we argue, through a simple theoretical model, that such correlations arise from selection pressures for efficient learning of behavior sequences. We define intelligence operationally as...

Humans have genetically based unique abilities making complex culture possible; an assemblage of traits which we term "cultural capacity". The age of this capacity has for long been subject to controversy. We apply phylogenetic principles to date this capacity, integrating evidence from archaeology, genetics, paleoanthropology, and linguistics. We...

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Det kan framstå som nästan mirakulöst att det existerar någonting så intrikat och komplicerat i universum som en blomma eller en katt. Hur kan sådan komplexitet uppkomma och fungera? Från att ha varit ett av bevisen för en guds existens har vi nu tack vare Charles Darwin och forskarna i hans efterföljd en godvetenskaplig förståelse fö r hur det gåt...

We investigated whether a sexual preference for smoking can be related to past experiences of parental smoking during childhood, as predicted by the theory of sexual imprinting, but also by sexual conditioning theory. In a sample of over 4000 respondents to five Internet surveys on sexual preferences, we found that parental smoking correlates with...

Because the failure to escape a predator causes death to a prey animal, and thus excludes opportunities to reproduce in the future, predation is a major selective force in nature (e.g., Lima and Dill 1990; Dawkins and Krebs 1979). To evade attacks from an array of different predators successfully is thus of key importance for all ­potential prey or...

What determines the number of cultural traits present in chimpanzee (Pan troglodytes) communities is poorly understood. In humans, theoretical models suggest that the frequency of cultural traits can be predicted by population size. In chimpanzees, however, females seem to have a particularly important role as cultural carriers. Female chimpanzees...

1. The shape of the function linking predator-attack success rate with distance to predator-concealing cover, or prey refuge, will affect population dynamics, distribution patterns and community trophic structure. Theory predicts that predator-attack success should decline exponentially with distance from predator-concealing cover, resulting in a t...

Variation in antipredation behavior should translate into variation in survival. Effective general defenses, such as predator avoidance, decrease the likelihood that an individual is attacked and should therefore reduce selection on behaviors that enhance probability of escape on attack. Escape behaviors become important if animals cannot avoid att...

Bird and Emery (1) showed that rooks (Corvus frugilegus) can learn to manufacture and use tools to obtain food. They suggest that these behaviors emerge through insight and the authors touch upon a fundamental question in the study of animal intelligence: How can insight learning be separated from shaping?

Animals gain antipredation benefits from being in larger groups through increased probability of predator detection, dilution of individual risk of being attacked and confusion of predators during attack. A further benefit is that individuals in larger groups can decrease the amount of time they spend being vigilant, while maintaining a high probab...

Recent studies have shown that migratory thrush nightingales (Luscinia luscinia) experimentally treated with multiple changes of the magnetic field simulating a journey to their target stopover area in northern Egypt, increased fuel deposition as expected in preparation to cross the Sahara desert. To investigate the significance of food intake on t...

In theory, survival rates and consequent population status might be predictable from instantaneous behavioural measures of how animals prioritize foraging vs. avoiding predation. We show, for the 30 most common small bird species ringed in the UK, that one quarter respond to higher predation risk as if it is mass-dependent and lose mass. Half respo...

The use of metal and colour-rings or bands as a means of measuring survival, movements and behaviour in birds is universal and fundamental to testing ecological and evolutionary theories. The practice rests on the largely untested assumption that the rings do not affect survival. However this assumption may not hold for several reasons, for example...

The effect of competition for a limiting resource on the population dynamics of competitors is usually assumed to operate directly through starvation, yet may also affect survival indirectly through behaviourally mediated effects that affect risk of predation. Thus, competition can affect more than two trophic levels, and we aim here to provide an...

Predation and predation risk have recently been shown to have profound effects on bird migration, but we still know relatively little about how birds respond to predation risk en route and how this is translated into fundamental aspects of optimal migration. Here, we make the case that to understand the fitness consequences of foraging and anti-pre...

Butterflies that hibernate exhibit particularly efficient defence against predation. A first line of defence is crypsis, and most hibernating butterflies are leaf mimics. When discovered, some species have a second line of defence; the peacock, I. io, when attacked by a predator flicks its wings open exposing large eyespots and performs an intimida...

It has been suggested that bilateral symmetry may impose a cost for animals relying on camouflage because symmetric color patterns might increase the risk of detection. We tested the effect of symmetry on crypsis, carrying out a predation experiment with great tits (Parus major) and black-and-white--patterned, artificial prey items and background....

Any animal whose form or behavior facilitates the avoidance of predators or escape when attacked by predators will have a greater probability of surviving to breed and therefore greater probability of producing offspring (i.e., fitness). Although in theory the fitness consequences of any antipredation behavior can simply be measured by the resultan...

Long-lived butterflies that hibernate as adults are expected to have well-developed antipredation devices as a result of their long exposure to natural enemies. The peacock butterfly, Inachis io, for instance, is a cryptic leaf mimic when resting, but shifts to active defence when disturbed, performing a repeated sequence of movements exposing majo...

Cryptic prey coloration typically bears a resemblance to the habitat the prey uses. It has been suggested that coloration which visually matches a random sample of the background maximizes background matching. We studied this previously untested hypothesis, as well as another, little studied principle of concealment, disruptive coloration, and whet...

To investigate flight ability in captive Zebra Finches during reproduction we compared change in escape take‐off ability and wing load of reproducing females with their mates and non‐reproducing females when attacked by a model raptor. Initially females had 18% higher wing load than males. Non‐reproducing females and females that had started egg‐la...

To maximise survival during foraging animals must decide when and for how long forag-ing should be interrupted in order to avoid predators. Previous experiments have shown that birds that hear other individuals'alarm calls resume feeding later than those that see a flying predator. However, the responses of prey animals to enemies are highly contex...

Migrating birds must accumulate fuel during their journeys and this fuel load should incur an increased risk of predation. Migratory fuelling should increase individual mass-dependent predation risk for two reasons. First, acquisition costs are connected to the increased time a bird must spend foraging to accumulate the fuel loads and the reduced p...

To cope with fluctuating environments animals have evolved reversible phenotypic flexi-bility. Some birds demonstrate this phenomenon by changing mass and flight muscle ac-cording to changes in wing loading. During moult, birds suffer from reduced wing area because feathers are shed and replaced, resulting in a wing loading increase. Moult is rathe...

To detect threats and reduce predation risk prey animals need to be alert. Early predator detection and rapid anti-predatory action increase the likelihood of survival. We investigated how foraging affects predator detection and time to take-off in blue tits (Parus caeruleus) by subjecting them to a simulated raptor attack. To investigate the impac...

The dilution effect as an antipredation behaviour is the main theoretical reason for grouping in animals and states that all individuals in a group have an equal risk of being predated if equally spaced from each other and the predator. Stalking predators, however, increase their chance of attack success by preferentially targeting nonvigilant indi...

When birds are attacked by aerial predators they should benefit by adjusting their escape to the prevailing attack situation. One important factor likely to affect escape decisions of prey, to our knowledge not previously studied, is the distance at which the attacking predator is detected. We investigated if great tits Parus major and blue tits P....

Bird migration requires high energy expenditure, and long-distance migrants accumulate fat for use as fuel during stopovers throughout their journey. Recent studies have shown that long-distance migratory birds, besides accumulating fat for use as fuel, also show adaptive phenotypic flexibility in several organs during migration. The migratory rout...

Bird mortality is heavily affected by birds of prey. Under attack, take-off is crucial for survival and even minor mistakes in initial escape response can have devastating consequences. Birds may respond differently depending on the character of the predator's attack and these split-second decisions were studied using a model merlin (Falco columbar...

Studies of naturally predator-naive adult birds (finches on predator-free islands)and birds experimentally hand reared in isolation from predators indicate that birds can recognise predators innately; that is, birds show anti-predator behaviour without former experience of predators. To reduce predation risk efficiently during the vulnerable fledgl...

Impaired predator evasion in birds as a cost in different life‐history periods has received increasing attention in the last decade. Evasive abilities in birds have been found to be detrimentally affected by migratory fuel load, reproduction and moult. These results suggest that during these periods of their lives birds suffer from increased predat...

Long stretches of sea and desert often interrupt the migration routes of small songbirds, whose fat reserves must be restored before these can be crossed as they provide no opportunity for refuelling. To investigate whether magnetic cues might enable inexperienced migratory birds to recognize a region where they need to replenish their body fat, we...

Environmental changes are responsible for the evolution of flexible physiology and the extent of phenotypic plasticity in the regulation of birds' organ size has not been appreciated until recently. Rapid reversible physiological changes during different life-history stages are virtually only known from long-distance migrants, and few studies have...

The period between independence and the early phase of autumn migration is a difficult period for juvenile birds and a relatively poorly known part of the annual cycle. We present data on post-juvenile moult, fuel deposition and speed of early autumn migration of "red-spotted" Blucthroats Luscinia s. svecica, studied during 14 years (1984-1997) in...

The secretive, endemic Short-legged Ground-Roller Brachypteracias leptosomus was studied from October 1996 to February 1997 on the Masoala Peninsula, northeast Madagascar. Several vocalizations were associated with contact, courtship feeding and food solicitation. One study pair ranged within an area of 19.1 ha and spent 90% of their time together....

Recent studies have shown that large fuel loads in small birds impair flying ability. This is the first study to show how migratory fuel load affects flying ability, such as velocity and height gained at take-off in a predator escape situation, in a medium-distance migrant, and whether they adjust their take-off according to predator attack angle....