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Fixed action pattern

A fixed action pattern (FAP) is an instinctive, highly stereotyped sequence of behaviors in animals that is innate, unlearned, and triggered by a specific external stimulus known as a sign stimulus or releaser, proceeding to completion once initiated regardless of changes in the environment or removal of the stimulus.^[1]^[2] These patterns are species-specific and serve essential functions in survival, reproduction, and social interaction, forming a core concept in ethology, the biological study of animal behavior.^[1]^[3] The concept of FAPs emerged in the 1930s through the pioneering work of Austrian ethologist Konrad Lorenz and Dutch ethologist Niko Tinbergen, who described them as rigid, invariant motor programs hard-wired into the nervous system and executed via central pattern generators.^[1]^[2] Triggering occurs through innate releasing mechanisms (IRMs), neural filters that recognize and process the sign stimulus to initiate the behavior, often independently of learning or external feedback.^[2] FAPs differ from simple reflexes by their complexity and coordinated sequence, with traits such as predictability, automaticity, and lack of modification across individuals of the same species.^[1]^[4] Classic examples illustrate FAPs' precision and evolutionary significance: in greylag geese, an egg displaced from the nest prompts a fixed rolling motion with the beak to retrieve it, halting only upon success; similarly, male three-spined stickleback fish display aggressive postures toward any red object, mistaking it for a rival during breeding.^[1]^[3] In insects like crickets, FAPs manifest as stridulation songs for mate attraction, coordinated by thoracic ganglia.^[2] Although early models emphasized their "fixed" nature, modern ethological and neuroethological research acknowledges variability, referring to them as modal action patterns (MAPs) to account for individual differences and contextual modulation while retaining their innate core.^[4]^[1] FAPs continue to inform studies in behavioral genetics, evolutionary biology, and information theory, where their stereotypy is quantified to distinguish innate from learned actions.^[4]^[1]

Definition and History

Core Definition

A fixed action pattern (FAP) is a species-specific, innate sequence of behaviors that is triggered by a particular stimulus and executed in a highly stereotyped manner until completion, regardless of environmental changes once initiated.^[5] These patterns represent genetically programmed motor coordinations that are characteristic of a species and serve adaptive functions in survival and reproduction.^[5] Unlike variable or flexible responses, FAPs exhibit a fixed form and sequence, making them reliable indicators of phylogenetic relationships among animals.^[5] The core components of an FAP include the releasing stimulus, also known as the sign stimulus, which is a specific external cue that initiates the behavior; the innate releasing mechanism (IRM), a neural filter that detects and responds to this stimulus without prior learning; and the fixed motor pattern itself, which unfolds as a coordinated, centrally driven sequence.^[5] The IRM acts as a selective physiological detector, ensuring that only relevant, biologically significant stimuli trigger the response.^[6] Once activated, the FAP proceeds endogenously to a consummatory act that reduces the underlying drive, often following an "all-or-nothing" principle where the behavior runs its full course even if the initial stimulus is withdrawn.^[5] FAPs are fundamentally distinct from learned behaviors, which develop through experience, trial, or imitation, as they are hardwired and resistant to modification by individual learning or environmental feedback.^[6] They also differ from simple reflexes, which are brief and stimulus-bound, whereas FAPs involve complex, prolonged sequences driven by internal motivation rather than direct sensory input.^[5] This innate, unmodifiable nature ensures their consistency across individuals of the same species.^[5] The concept of FAPs was originally conceptualized in the early 20th century by ethologist Oskar Heinroth, who introduced the term "arteigene Triebhandlung" to describe these species-characteristic drive actions.^[5] His work laid the groundwork for later refinements by Konrad Lorenz and Niko Tinbergen, who formalized the English term "fixed action pattern" in their ethological studies of the 1930s–1950s, establishing FAPs as a cornerstone of ethological theory.^[6]^[7]

Historical Development

The concept of fixed action patterns emerged from early 20th-century observations of instinctive behaviors in birds, pioneered by German ornithologist Oskar Heinroth. In the 1910s, Heinroth conducted detailed comparative studies of waterfowl, documenting species-specific behavioral sequences such as mating displays and molting cycles that appeared innate and stereotyped, linking them to ecological adaptations like mate attraction and communication.^[7] His 1911 publication emphasized these behaviors' reliability across individuals, laying foundational groundwork for ethology by highlighting their role in taxonomic classification and functional significance, though he did not yet formalize the term "fixed action pattern."^[8] In the 1930s and 1940s, Austrian biologist Konrad Lorenz built upon Heinroth's observations, refining the notion of innate motor patterns as genetically determined "fixed action patterns" (FAPs) that unfold in rigid sequences once triggered. Collaborating with Dutch ethologist Niko Tinbergen, Lorenz developed the hydraulic model of motivation around 1937–1950, conceptualizing FAPs as driven by accumulating "action-specific energy" in a reservoir, released through innate releasing mechanisms in response to environmental stimuli; this model explained phenomena like vacuum activities when energy built without release.^[9] Tinbergen complemented this theoretical framework with rigorous field experiments in the 1930s–1950s, validating FAPs through controlled manipulations, such as studies on egg-rolling in greylag geese (1938) and gaping responses in thrushes (1939), demonstrating the precision of stimulus-response thresholds and hierarchical organization of behaviors.^[7] Their joint work, culminating in Tinbergen's 1951 book The Study of Instinct, established FAPs as a core ethological principle, integrating observation with experimental testing.^[7] Following World War II, the 1950s saw FAP research integrate into broader ethology and neurobiology, with Lorenz and Tinbergen's contributions gaining international recognition. In 1973, they shared the Nobel Prize in Physiology or Medicine with Karl von Frisch for discoveries concerning "the organization and elicitation of individual and social behavior patterns," underscoring FAPs' role in understanding innate behavioral coordination.^[10] By the 1980s, while FAPs retained relevance in behavioral ecology for explaining adaptive responses, critiques challenged the emphasis on innateness, arguing that behaviors exhibit greater flexibility and environmental influence than Lorenz's rigid model suggested. Influential debates, sparked by comparative psychologists like Daniel Lehrman in the 1970s and continuing into the 1980s, prompted a shift toward integrating learning and plasticity, with behavioral ecology adopting quantitative models to assess adaptive variability over strict instinctual determinism.^[11] This evolution refined FAPs as part of a dynamic framework, maintaining their utility in studying species-typical traits amid ongoing neurobiological insights.^[11]

Key Characteristics

Stereotyped Nature

Fixed action patterns (FAPs) are characterized by their highly stereotyped execution, consisting of a rigid, unvarying sequence of movements that follows a predictable order once initiated. This stereotypy manifests as a "ballistic" performance, where the behavior proceeds to completion with minimal modification, even in the face of environmental disruptions or interruptions, ensuring a consistent motor output across occurrences.^[12] Such invariance underscores the pre-programmed nature of FAPs, distinguishing them from more flexible, learned behaviors, though modern views recognize some limited individual and contextual variability.^[2]^[4] Observational evidence for this stereotyped nature comes from ethological studies demonstrating remarkable consistency in the timing, posture, and motor patterns of FAPs when performed repeatedly by individuals within the same species. Across multiple trials and contexts, these behaviors exhibit low variability, with quantitative metrics such as movement trajectories and durations showing minimal deviations, often assessed through techniques like standard deviation analyses that highlight their reliability.^[12] Pioneering work by Konrad Lorenz and Niko Tinbergen emphasized this uniformity, describing FAPs as innate, coordinated patterns that remain stable regardless of external influences.^[2] At the neural level, the stereotyped execution of FAPs is underpinned by central pattern generators (CPGs), neural circuits typically located in the spinal cord or brainstem that autonomously produce rhythmic or sequential motor outputs without requiring continuous sensory input. These CPGs enable the automatic, hard-wired coordination of muscle activity, contributing to the ballistic and resistant quality of the behavior.^[12] Ethological measurements of FAP stereotypy often involve high-resolution techniques, such as electromyography or kinematic tracking, to assess variability in key parameters like amplitude and phase.^[2]

Innate and Species-Specific Traits

Fixed action patterns (FAPs) represent innate behaviors that manifest without any prior learning or experience, emerging spontaneously upon initial exposure to a sign stimulus. This unlearned quality is particularly evident in hatchlings and juveniles, where the full sequence of the pattern is performed correctly from the outset. For example, newly hatched greylag goose chicks instinctively roll displaced eggs back to the nest using a precise head movement, even if the egg is removed midway through the action, demonstrating the pattern's autonomous completion.^[13] Such observations in naive animals, including incubator-reared birds isolated from conspecifics, confirm that FAPs are pre-programmed and require no observational learning or practice to execute species-typical responses.^[14] The species-specific nature of FAPs underscores their adaptation to the unique ecological niches of individual taxa, rendering them non-transferable across different species. These patterns vary distinctly to suit environmental demands, such as the courtship zig-zag swimming and red belly display in male three-spined stickleback fish, which signal readiness to mates in shallow freshwater habitats, in contrast to the song repertoires and aerial chases of passerine birds tailored for territorial defense in arboreal settings.^[13] This specificity ensures that each FAP aligns with the sensory and motor capabilities evolved within a species, preventing maladaptive cross-species mimicry.^[2] Genetic underpinnings of FAPs are rooted in their congenital preformation, with heritability evidenced by the consistent transmission of these behaviors across generations in breeding populations. In model organisms like orthopteran insects, stridulatory FAPs—such as the rhythmic wing movements producing species-specific calls—are genetically determined and appear without environmental modification, as shown in isolated rearing studies that eliminate learning influences.^[2] Ethological experiments, including selective breeding in birds and fish, further demonstrate high heritability for motor components of FAPs, where offspring reliably exhibit parental patterns despite varied rearing conditions.^[14] During developmental ontogeny, FAPs mature in a predictable sequence aligned with an animal's growth phases, transitioning from rudimentary expressions in juveniles to refined adult forms. Similarly, in birds, innate pecking or retrieval behaviors are operational at hatching but gain precision through physiological development, ensuring readiness for ecological roles at independence.^[14] This ontogenetic progression highlights the intrinsic timing of FAP emergence, contributing to their stereotyped reliability in behavioral execution.

Triggering Mechanisms

Sign Stimuli

Sign stimuli, also known as releasers, are specific environmental cues—often minimal and simple signals such as shapes, colors, or movements—that activate the innate releasing mechanism (IRM), a neural template in the central nervous system that initiates fixed action patterns (FAPs).^[2] These stimuli were first conceptualized by Konrad Lorenz as "Angeborenes Auslösendes Schema," emphasizing their role in triggering innate behaviors without requiring learning or complex processing.^[2] In essence, sign stimuli serve as precise triggers tuned by evolution to ensure reliable responses in species-typical contexts. The function of sign stimuli operates analogously to a key fitting a lock within the nervous system, where the stimulus matches the IRM's sensory filter, thereby disinhibiting and releasing the coordinated motor program of the FAP.^[2] This mechanism bypasses higher cognitive evaluation, allowing for rapid, automatic execution of essential survival behaviors like aggression or courtship, as refined in Niko Tinbergen's framework of the IRM.^[2] The simplicity of these cues—focusing on salient features rather than full contextual details—facilitates efficient activation while minimizing false positives in variable natural environments. Experimental identification of sign stimuli typically involves presenting isolated or artificial models to subjects to pinpoint the minimal effective components. For example, Tinbergen's 1951 studies on three-spined sticklebacks used crude dummy models to demonstrate that a red underside alone elicited intense aggressive FAPs from territorial males, whereas models lacking this color feature provoked little response, isolating the red coloration as the critical releaser.^[15] Such methods highlight the precision required, as only specific stimulus configurations successfully engage the IRM. The specificity of sign stimuli is evolutionarily honed, with a narrow range of effective cues ensuring activation occurs reliably under natural conditions where relevant signals are likely to appear.^[16] This tuning prevents over-response to irrelevant inputs, promoting adaptive efficiency in behaviors critical for reproduction and survival, as evidenced in Tinbergen's ethological experiments that underscored the IRM's selective responsiveness.^[2]

Supernormal Stimuli

Supernormal stimuli represent exaggerated versions of natural sign stimuli that elicit a more intense and prolonged fixed action pattern response than the typical environmental cues for which the behavior evolved.^[17] These artificial or amplified signals exploit the innate releasing mechanisms underlying fixed action patterns, often leading animals to prioritize the supernormal cue over more adaptive natural ones. The concept was first systematically explored by ethologist Niko Tinbergen, who demonstrated how such stimuli can override normal behavioral thresholds, resulting in maladaptive overreactions.^[18] A classic example involves the egg-incubation behavior in Eurasian oystercatchers (Haematopus ostralegus). Normally, these birds lay and incubate clutches of three eggs, but when presented with artificial clutches of five eggs or oversized dummy eggs, females preferentially incubated the larger sets or the biggest individual egg, even if it was too large to cover properly or balance on their nest.^[19] This exaggerated preference hijacks the fixed action pattern of egg retrieval and brooding, triggered by visual cues like size and shape, illustrating how supernormal stimuli amplify innate responses beyond ecological utility.^[19] In herring gulls (Larus argentatus), supernormal stimuli affect the food-begging fixed action pattern of chicks. Tinbergen and colleague A.C. Perdeck found that newly hatched chicks pecked more vigorously at cardboard models featuring elongated bills with high-contrast red spots and white bands compared to the natural parental beak, which has a shorter bill and subtler markings. The chicks even ignored realistic three-dimensional models in favor of these simplified, exaggerated versions, showing how supernormal cues—such as increased length, contrast, or motion—intensify the pecking response, a stereotyped sequence of head movements and bill snaps. These phenomena highlight the rigidity of fixed action patterns, where evolutionary tuning to average natural stimuli leaves behaviors vulnerable to extreme exaggerations. While adaptive for detecting slightly superior natural variants (e.g., larger eggs signaling better nutrition), supernormal stimuli reveal potential evolutionary pitfalls, as animals may expend energy on non-viable objects.^[19] Tinbergen's experiments underscore the hierarchical nature of stimulus-response systems, with supernormal cues often dominating due to their peak alignment with innate perceptual biases.^[19]

Illustrative Examples

Stickleback Fish Mating Ritual

The three-spined stickleback (Gasterosteus aculeatus) provides a seminal case study of fixed action patterns (FAPs) in male aggressive and courtship behaviors, extensively investigated by ethologist Nikolaas Tinbergen in the 1950s. Inhabiting shallow freshwater and coastal habitats across the Northern Hemisphere, these small fish breed in spring, where males establish and defend territories to facilitate reproduction. During this period, breeding males undergo physiological changes, developing a bright red nuptial coloration on their throat and belly, which serves as a key visual cue in social interactions.^[20] The aggressive FAP in territorial males is triggered by the sign stimulus of a red belly on an intruding rival, initiating a stereotyped sequence designed for territory defense. Upon detection, the male approaches the intruder with rapid zigzag swimming—alternating straight rushes toward the stimulus and sharp lateral turns—followed by a head-up threat display, in which the male erects its dorsal spines, tilts its head upward to expose its own red throat, and holds a rigid posture. If the rival persists, the sequence escalates to direct charges, chases, and biting attacks, often culminating in the intruder's flight and reinforcement of nest-building activities to secure the territory. Tinbergen's experiments in the early 1950s used crude dummy models to isolate this sign stimulus: a realistic gray model elicited minimal response, whereas adding a red underside provoked the full aggressive motor pattern, even in naive males, demonstrating the innate and specific nature of the trigger.^[21] In parallel, the courtship FAP targets receptive females and runs as a complete, irreversible sequence from approach to fertilization. The primary sign stimulus is the gravid female's swollen abdomen, signaling ripeness for spawning. Encountering this, the male executes zigzag swimming to orient and attract her, transitioning to straight-lead swimming toward the nest—a tubular structure of plant material glued with kidney secretions that the male constructs in advance. At the nest entrance, the male performs a head-down display, tilting to point at the opening, then quivers rapidly against the female's side to stimulate egg release. The female enters and deposits eggs, after which the male fertilizes them by releasing milt, completing the pattern and shifting to parental care. Tinbergen's dummy model tests confirmed the swollen abdomen's potency: models with exaggerated ventral swelling elicited vigorous courtship, including the full zigzag and leading behaviors, underscoring the FAP's release by minimal, species-specific cues.^[20] These aggressive and courtship sequences in the stickleback exemplify FAPs' role in efficient territory defense and reproductive success within resource-limited freshwater environments, where rapid, predictable responses minimize energy expenditure and maximize mating opportunities.^[21]

Greylag Goose Egg Retrieval

One of the most iconic examples of a fixed action pattern is the egg-retrieval behavior observed in Greylag geese (Anser anser) by Konrad Lorenz and Niko Tinbergen. When an egg becomes displaced from the nest, the female goose responds with a highly stereotyped sequence: she extends her neck downward toward the egg, positions her beak behind it, and uses a combination of forward and lateral head movements while walking backward to roll the egg back to the nest.^[1]^[22] This behavior is triggered specifically by the sign stimulus of an egg visible outside the nest boundaries, initiating the full motor pattern without requiring additional cues. Remarkably, the sequence persists to completion even if the egg is removed mid-process; the goose continues the rolling motion with her beak and body as if the egg remains present, often "tucking" an imaginary egg into the nest upon arrival. This persistence highlights the pattern's ballistic nature, driven by central neural coordination rather than ongoing sensory feedback.^[1]^[23]^[22] The egg-retrieval action is innate, executed identically by naive young geese that have never observed or practiced it, demonstrating its genetic basis and independence from learning. Lorenz's detailed observations underscored this innateness, noting that hand-reared goslings performed the full sequence flawlessly upon first encountering a displaced egg. Such experimental evidence from Lorenz and Tinbergen's work established the pattern as a model for understanding centrally coordinated instinctive behaviors in vertebrates.^[1]^[23]^[22]

Additional Animal Cases

In insects, female silk moths (Bombyx mori) exhibit a fixed orientation response to the male-released bombykol pheromone, triggering a stereotyped upwind flight and landing sequence toward the source, even with naive individuals, demonstrating innate chemotaxis as a FAP for mate location.^[1] Among mammals, laboratory rats (Rattus norvegicus) produce ultrasonic vocalizations (50-60 kHz) as pups when isolated from the mother, eliciting a stereotyped retrieval and nursing response in the dam, an innate pattern conserved across rodents for survival.^[2] In amphibians, male túngara frogs (Engystomops pustulosus) produce croaking calls as a fixed vocalization pattern to attract females, consisting of a whine followed by optional chucks, triggered by auditory cues from rival males or environmental conditions like humidity.^[24] This advertisement call is species-specific and highly stereotyped, ensuring effective mate location in choruses despite acoustic interference.^[25] Fixed action patterns appear across phyla, particularly in survival-related functions such as foraging and alarm responses, where they provide rapid, reliable behaviors adapted to ecological pressures from insects' recruitment dances to mammals' threat displays.^[2] In reptiles and amphibians, similar innate sequences support predator evasion or territory defense, underscoring the evolutionary conservation of these mechanisms for essential life processes.^[26]

Evolutionary Perspectives

Adaptive Benefits

Fixed action patterns (FAPs) provide significant adaptive advantages by enabling animals to respond swiftly and effectively to environmental challenges, thereby enhancing overall fitness. Their stereotyped nature allows for immediate execution without the need for deliberation or learning, which is crucial in high-stakes situations such as predator avoidance or resource acquisition. This efficiency minimizes the time and cognitive resources required, conserving energy that can be allocated to other survival needs. These patterns have evolved through natural selection to promote survival and reproduction in ancestral environments.^[27] The innate quality of FAPs ensures reliability, particularly for inexperienced individuals like juveniles or those under stress, where learned behaviors might fail. By being genetically programmed, these patterns perform consistently across individuals within a species, reducing variability that could lead to suboptimal outcomes in critical contexts. For instance, in threat responses, the predictable activation via sign stimuli guarantees a coordinated defense without hesitation. This dependability is a cornerstone of ethological theory, as articulated in foundational work on instinctive behaviors. In the realm of reproduction, FAPs play a pivotal role in promoting mating success through synchronized displays that signal readiness and species identity. Courtship rituals, such as those observed in various birds and fish, facilitate partner attraction and reduce interspecies mating errors, directly contributing to gene propagation. These patterns evolve to align with selective pressures, ensuring that reproductive efforts yield higher offspring survival rates. Furthermore, FAPs exhibit ecological fitting by being finely tuned to the specific sensory and environmental cues prevalent in a species' habitat, optimizing responses to local conditions. This adaptation minimizes misdirected actions, such as inappropriate aggression or foraging, and enhances integration into ecological niches.

Potential Drawbacks

The rigidity inherent in fixed action patterns (FAPs) can lead to maladaptive outcomes, as these behaviors are typically performed to completion once initiated, regardless of intervening changes in circumstances. This "ballistic" nature means that animals may expend effort on irrelevant or counterproductive actions, such as a greylag goose continuing its egg-retrieval sequence by circling and pushing even after the egg has rolled away or been removed from the nest site. In such cases, the central nervous coordination driving the FAP prevents mid-sequence adaptation, resulting in unnecessary physical exertion without achieving the intended goal. In environments that deviate from ancestral conditions, FAPs tuned to specific ecological niches may fail to confer benefits and instead impose fitness costs. For instance, urban habitats introduce novel stimuli and risks, such as traffic or pollutants, where instinctive sequences optimized for natural settings may expose animals to heightened predation or injury without yielding survival advantages. Urban animals often require behavioral plasticity to adapt to these rapid changes, and a lack of flexibility in instinctive behaviors can contribute to challenges in survival and reproduction.^[28] The execution of FAPs also carries significant energetic demands, as these complex, stereotyped sequences require coordinated muscular activity and metabolic investment, even when triggered inappropriately. According to Lorenz's hydraulic model of motivation, accumulated action-specific energy propels the full performance of an FAP once released, depleting reserves that could otherwise support essential activities like foraging or predator avoidance. In altered ecosystems, such as fragmented habitats where sign stimuli are frequent but unreliable, repeated unnecessary activations of FAPs can accelerate energy exhaustion, contributing to lower overall fitness in affected populations.

Exploitation in Brood Parasitism

Brood parasites exploit the fixed nature of host fixed action patterns (FAPs) by mimicking the sign stimuli that trigger innate parental care behaviors, such as egg incubation and chick provisioning, thereby inducing hosts to invest resources in raising unrelated offspring.^[29] In avian systems, this manipulation begins at the egg stage, where parasites produce eggs that closely resemble those of their hosts in color, pattern, and size to evade the host's egg-rejection FAP, which is elicited by foreign appearances. For instance, the common cuckoo (Cuculus canorus) lays eggs that mimic the blue-green speckled pattern of the reed warbler (Acrocephalus scirpaceus), a frequent host, reducing the likelihood of recognition and ejection.^[30] At the chick stage, brood parasites further hijack host FAPs through behavioral and morphological mimicry that amplifies begging signals, exploiting the host's instinctive response to feed wide-gaped, vocalizing nestlings. The common cuckoo chick, upon hatching, rapidly ejects host eggs or chicks and then employs exaggerated begging calls and gapes with bright yellow flanges—often more conspicuous than those of host young—to solicit food from reed warbler parents, who continue provisioning despite the size disparity.^[29] Similarly, the brown-headed cowbird (Molothrus ater) in North American songbird hosts uses intense, repetitive begging displays that mimic a supernormal brood stimulus, prompting hosts like the prothonotary warbler (Protonotaria citrea) to allocate disproportionate feeding efforts.^[30] This exploitation has fueled an evolutionary arms race, where hosts develop refined sensory discrimination to counter parasitic mimicry, while parasites evolve tighter tuning to host-specific cues for greater success. Reed warblers, for example, have been observed to reject cuckoo eggs in populations with high parasitism rates, selecting for cuckoo gentes (host-specific races) with increasingly precise egg mimicry.^[30] Consequently, successful parasitism enhances parasite fitness by offloading all parental investment onto hosts, often resulting in near-complete fledging success for the parasite at the expense of host reproductive output, as a single cuckoo chick can consume resources equivalent to an entire host brood.^[29]

Variations and Exceptions

Threshold Reduction Effects

In Konrad Lorenz's hydraulic model of motivation, action-specific energy accumulates within a species-specific reservoir for each fixed action pattern (FAP), building up continuously during periods of deprivation and lowering the response threshold required to trigger the behavior.^[5] This endogenous drive, termed action-specific potential (ASP), intensifies motivational states, making the innate releasing mechanism more sensitive to even weak or absent stimuli over time.^[5] As deprivation prolongs, the dropping threshold can result in partial or complete spontaneous expression of FAPs without an external releasing stimulus, allowing the accumulated energy to dissipate through innate motor patterns.^[31] For instance, in birds deprived of breeding opportunities, such as canaries held in captivity without nesting materials during spring, individuals may initiate full nest-building sequences, including gathering and arranging imaginary or inadequate objects, to release the pent-up ASP.^[5] This phenomenon underscores how FAPs, though stereotyped, adapt to internal pressures by enabling autonomous outbursts that maintain behavioral readiness.^[31] Experimental studies on birds have demonstrated this threshold reduction empirically, showing increased responsiveness to suboptimal stimuli following extended deprivation periods. In investigations of canary nest-building, Robert Hinde observed that prolonged isolation led to spontaneous initiation of FAP components, such as material manipulation and cup-molding, with thresholds for triggering behaviors fluctuating based on time since last performance.^[5] Similarly, research on red-backed shrikes (Lanius spp.) revealed that hunger deprivation lowered the threshold for prey-impaling FAPs, prompting birds to skewer non-prey items like twigs when real food was unavailable, confirming the role of ASP accumulation in heightening behavioral excitability.^[5] The biological basis of these threshold effects involves central nervous system changes in motivational states, where accumulated ASP generates endogenous impulses that enhance neural excitability independent of external input.^[5] This may arise from neurotransmitter accumulation altering synaptic responsivity in relevant motor circuits, as seen in model systems where deprivation sensitizes neural pathways to lower activation thresholds.^[31] Such mechanisms ensure that FAPs remain phylogenetically programmed responses attuned to survival needs, with deprivation acting as a teleonomic regulator of instinctive readiness.^[5]

Vacuum and Displacement Activities

Vacuum activities represent instances where an animal performs elements of a fixed action pattern (FAP) in the complete absence of the usual releasing stimulus, driven by a buildup of internal motivation. These behaviors occur when action-specific potential (ASP), the endogenous excitability associated with a particular instinct, accumulates to such an extent that the response threshold is drastically lowered, sometimes approaching zero. As a result, the animal may execute partial or complete motor patterns spontaneously, often with substitute objects or no object at all. A classic example is observed in greylag geese, where individuals engage in egg-rolling motions without an actual egg present, grasping and maneuvering imaginary objects as if retrieving one.^[5] This phenomenon underscores the innate, centrally coordinated nature of FAPs, which can be triggered endogenously by neural processes independent of external afferent input.^[5] Displacement activities, in contrast, emerge during situations of motivational conflict, where an animal experiences simultaneous drives toward incompatible behaviors, such as aggression and fear. In these cases, the primary FAPs are blocked or inhibited, leading to the insertion of an irrelevant, unrelated behavior derived from another instinctual system. For instance, in male three-spined sticklebacks (Gasterosteus aculeatus), when the fighting drive is obstructed—such as during territorial disputes near a boundary—individuals perform displacement digging, a nest-building activity that appears out of context and unrelated to the ongoing conflict.^[32] Similarly, during courtship interruptions, frustrated mating drives may redirect into pushing, fanning, or gluing behaviors, which serve as outlets for the pent-up energy.^[32] These activities are species-specific and innate, often lacking any immediate adaptive function in the given context but indicating underlying tension.^[32] The underlying mechanism for both vacuum and displacement activities is explained through Konrad Lorenz's hydraulic model of motivation, which conceptualizes ASP as a fluid-like energy that accumulates continuously within the central nervous system. In vacuum activities, prolonged absence of stimuli causes this energy to "dam up," resulting in overflow and spontaneous discharge of the FAP, potentially leading to hallucinatory-like responses where the animal reacts to internal impulses as if stimuli were present.^[5] For displacement activities, conflicting drives create mutual inhibition, neutralizing the primary actions and allowing the excess energy to "spark over" into a secondary, unrelated motor pattern, such as preening or feeding during agonistic encounters.^[5] This redirection prevents complete behavioral paralysis and provides a temporary outlet for excitation. In some cases, these behaviors may become ritualized over evolutionary time, acquiring signal functions, though many remain non-communicative.^[32] Such activities are commonly documented in both laboratory experiments and wild observations, serving as reliable indicators of an animal's internal motivational state. In captive settings, vacuum activities like nest-building in weaverbirds without materials or snapping motions in starlings at empty air highlight the persistence of instinctive readiness despite environmental deprivation.^[5] Displacement behaviors, meanwhile, frequently appear in natural conflicts, such as bill-shaking in terns torn between fleeing and incubating, revealing the dynamic interplay of drives.^[5] These manifestations not only demonstrate the robustness of FAPs under suboptimal conditions but also emphasize their role in revealing hidden aspects of animal psychology, as noted in ethological studies since the mid-20th century.^[5]

Graded and Flexible Responses

Fixed action patterns (FAPs) exhibit graded responses, where the intensity or vigor of the behavioral sequence varies proportionally with the strength of the sign stimulus or the animal's internal motivational state. For instance, in male three-spined sticklebacks (Gasterosteus aculeatus), the zigzag courtship display becomes faster and more pronounced when encountering a highly attractive female model compared to a less stimulating one, reflecting heightened arousal levels. Similarly, in crickets (Gryllus bimaculatus), the amplitude and rate of stridulation during calling songs increase with the proximity or responsiveness of a female receiver, modulated by sensory feedback to command neurons in the central nervous system.^[33] This gradation allows FAPs to adapt subtly to immediate contextual demands without altering the core sequence, as demonstrated in neurophysiological recordings of thoracic ganglia activity.^[33] Evidence of flexibility in FAPs emerged prominently in studies from the 1970s to the 2000s, revealing that these behaviors are not invariably rigid but can be modified by experience or environmental context, thus challenging the classical view of strict innateness. Schleidt's analysis of crowing in Japanese quail (Coturnix japonica) quantified intra- and inter-individual variability in the FAP, showing that while the pattern remains species-typical, its timing and amplitude adjust based on prior social exposure, with coefficients of variation indicating non-stereotyped elements.^[34] In sharks, such as the grey reef shark (Carcharhinus amblyrhynchos), agonistic displays like the hunch posture escalate in intensity—from mild stiffening to full pectoral fin depression and jaw gaping—in response to repeated intruder approaches, incorporating learned threat assessment from past encounters.^[35] These findings, including Barlow's concept of modal action patterns, highlighted how FAPs incorporate probabilistic variations, blending innate coordination with experiential tuning.^[36] Contemporary neuroethological perspectives propose hybrid models that integrate FAPs with learning mechanisms, portraying them as outputs of central pattern generators (CPGs) amenable to modulation by higher brain centers. In orthopterans, for example, the stereotyped wing movements for song production are driven by CPGs but fine-tuned through auditory feedback loops that incorporate contextual learning, such as adjusting chirp duration to optimize female attraction.^[33] This synthesis is evident in studies of acoustic communication, where innate releasing mechanisms interact with plastic neural circuits to allow adaptive flexibility without full relearning.^[33] Such models underscore FAPs as dynamic systems rather than immutable reflexes, with implications for bridging classical ethology—focused on instinctive rigidity—with cognitive behaviorism, which emphasizes environmental influence on behavioral expression.^[33]

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By studying “naive” animals (e.g. young birds born in an incubator), he was able to prove that these fixed action patterns appeared as reactions to key stimuli ...<|separator|>
[14]
The Distinction Between Innate and Acquired Characteristics
Aug 4, 2009 · Innate behavior was thought to be composed of identifiable units – 'fixed action patterns' – whose evolution could be studied like that of a ...
[15]
Ethological studies of sign stimuli and motivation - Fly Fishing Devon
Fixed Action Patterns. Lorenz considered that every species has a repertoire of stereotyped behaviours called Fixed Action Patterns. Lorenz considered that they ...
[16]
Fixed Action Pattern - an overview | ScienceDirect Topics
A fixed action pattern (FAP) is defined as an instinctive, indivisible behavioral sequence that, once initiated by a sign stimulus, will run to completion ...
[17]
Measuring Preference for Supernormal Over Natural Rewards
Oct 29, 2015 · Tinbergen (1948) coined the term “Supernormal Stimulus” upon finding that animals tend to exhibit heightened responses to exaggerated versions ...
[18]
https://www.jstor.org/stable/4157642
[19]
[PDF] Supernormal Stimuli and Responses
In this article, I describe the curious phenomenon of exagger- ated responses to supernormal stimuli in animals. These have.
[20]
The three-spined stickleback as a model for behavioural neuroscience
Mar 26, 2019 · The aim of the present study is to describe some innate behaviours of laboratory bred three-spined sticklebacks by using a battery of tests.Missing: spot | Show results with:spot
[21]
The Study of Instinct - Niko Tinbergen - Google Books
... sign stimuli social releasers species stimulus situation survival value territory three-spined stickleback Tinbergen tion tropisms visual wasp Weiss whole young ...
[22]
Taxis und Instinkthandlung in der Eirollbewegung der Graugans. I1
Aug 6, 2025 · An example of a fixed action pattern is the egg-rolling behavior of the graylag goose (Lorenz and Tinbergen, 1939) . When a female goose sees an ...
[23]
Simplification, Innateness, and the Absorption of Meaning from ...
The classic example of a FAP—egg rolling in the greylag goose (Anser anser)—also shows fusion. Upon seeing an egg placed by the side of its nest, the goose ...
[24]
Neuroethology of the Waggle Dance: How Followers Interact with ...
Oct 11, 2019 · Karl von Frisch discovered the meanings of the waggle dance and called the communication a “dance language.” Subsequent to this discovery, it ...
[25]
Learning in the honey bee waggle dance - SpringerLink
Jun 22, 2023 · Decades of work have gone into describing and studying how the waggle dance enables bees to find resources (von Frisch, 1967), and new work ...
[26]
[PDF] Lecture 29 Notes: Konrad Lorenz on learning - MIT OpenCourseWare
When the dog is shackled in an apparatus, the only aspect of the appetitive behavior it could show was salivation. Release it, and it shows begging behavior.
[27]
[PDF] Lecture 9 Notes: Lorenz on fundamentals of ethology
Cat must search much more than stalk, stalk more than pounce, etc. – The thresholds for stimuli to initiate searching (locomotion) are lowered the most quickly.Missing: sequence | Show results with:sequence
[28]
(PDF) Vocal communication in frogs - ResearchGate
Aug 7, 2025 · The robust nature of vocal communication in frogs has long attracted the attention of natural philosophers and their biologically inclined ...
[29]
Túngara frog call-timing decisions arise as internal rhythms interact ...
We investigated the factors influencing túngara frog call-timing responses to chorus-mates' calls on an interaction-by-interaction basis.
[30]
Motor control on the move: from insights in insects to general ...
This review discusses how the nervous system controls the complex body movements keeping animals up and running.<|control11|><|separator|>
[31]
https://us.sagepub.com/sites/default/files/upm-binaries/23196_Chapter_3.pdf
[32]
Animal cognition in an urbanised world - PMC - NIH
Urban environments often present animals with novel problems that differ from those encountered in their evolutionary past. To navigate these rapidly changing ...
[33]
Tricking Parents: A Review of Mechanisms and Signals of Host ...
Dec 2, 2021 · Here we review the literature pertaining to host manipulation by parasitic young, focusing on four non-mutually exclusive mechanisms.
[34]
Cuckoo adaptations: trickery and tuning - ZSL Publications - Wiley
Apr 18, 2011 · Cuckoo trickery involves adaptations to counter successive lines of host defence and includes: tricks for gaining access to host nests, egg ...Trickery: successive stages · Trickery: adaptation and lack... · Trickery versus tuning
[35]
The Modification of Instinctive Behavior
Tinbergen (1951) suggested that a central instinctive system (e.g., the reproductive instinct of stickleback fish) controls the occurrence of a number of ...
[36]
https://www.cambridge.org/core/journals/behavioral-and-brain-sciences/article/is-anything-fixed-in-an-action-pattern/BD9643894DD605CE4D5FACF5AE9FB8D8
[37]
[PDF] Konrad Lorenz 1956 Plays and Vacuum Activities In
When Monsieur le Professeur GRASSÉ invited me to talk about the relations between play and vacuum activity I said in my answering letter that I did not know ...
[38]
https://doi.org/10.1007/s00359-018-01311-3
[39]
Is anything fixed in an action pattern? | Behavioral and Brain Sciences
Feb 4, 2010 · Barlow, G. W. (1977) Modal action patterns. In: How animals ... 1969) Intracellular activity in cricket neurons during the generation of behavior ...