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. 2026 Jul 14;6(7):e70424. doi: 10.1002/cpz1.70424

Beyond Fear Learning in Isolation: Integration of Social Dimensions into Aversive Learning

Maria M Diehl 1,✉
PMCID: PMC13367259  PMID: 42446499

Abstract

Aversive learning paradigms in laboratory settings have led to valuable decades of research on the acquisition and expression of memories that produce fear and avoidance. However, the majority of these studies have focused on situations in which animals undergo aversive learning while they are in isolation. Although these studies have provided critical insights into the behavioral and neural processes guiding aversive learning, as well as translational insights into neuropsychiatric disorders, how social dimensions that are embedded in naturalistic aversive experiences influence these processes remains greatly understudied. This overview covers some of the aversive learning paradigms that have incorporated social dimensions to help us to understand how social information affects the learning of fear and avoidance. Key considerations and future directions for this line of research are also discussed. © 2026 Wiley Periodicals LLC.

Keywords: avoidance, fear, observational learning, rodents, social learning

INTRODUCTION

Imagine a small animal scampering through a field, when it suddenly detects a potential predatory threat: a shadow swiftly looming overhead, followed by an immediate unexpected sound. First, the mouse freezes, and then it quickly orients to the direction of the visual and auditory cues. If other conspecifics are nearby, the animal will do something that standard laboratory paradigms often fail to capture: look to other group members, evaluate their responses, and adjust its own responses accordingly. This naturalistic scenario captures an essential and well‐conserved process of nearly all mammalian life: the social regulation of fear.

The example above suggests that fear in response to threat is not always a solitary experience, especially for socially driven species. The standard laboratory paradigms that have defined our understanding of aversive learning (i.e., classical fear conditioning, extinction, active avoidance, etc.) have been largely developed around the isolated experience. This approach has been quite fruitful, giving us critical road maps for better understanding the neural circuits underlying the acquisition and expression of fear responses. Such paradigms have also led to many valuable insights into the etiology of multiple neuropsychiatric disorders and some of their treatments. Nevertheless, the translation from basic research to human neuropsychiatric disease remains a challenge. This might be due to the lack of social dimensions in aversive learning paradigms, which may bridge the translational gap between emotional processing and social cognition, given that anxiety disorders are often triggered by social cues or, conversely, can be treated through socially derived means (e.g., social coping). This overview covers some of the recent advances that researchers have accomplished to incorporate social dimensions into aversive learning. Some key variables are also identified that could reshape the impact of social information in aversive learning, along with discussion points worth considering for potential future research.

SOCIAL DIMENSIONS IN AVERSIVE LEARNING: WHAT HAS BEEN DONE AND WHAT HAS BEEN REVEALED?

Social dimensions that are often present during aversive experiences include the detection of particular behaviors and emotional signals from other conspecifics that are undergoing the same situation. For example, animals under immediate or potential danger emit specific vocalizations during threat detection. These vocalizations often alert other conspecifics in the group to pay attention and make the necessary behavioral adjustments to remain unharmed (e.g., freezing, escape, or avoidance). The following section provides a brief overview of several aversive learning paradigms that have been developed in a laboratory setting to incorporate social dimensions, such as aversive learning in groups, indirect aversive learning through social transmission of information, or associating a social stimulus with danger.

Fear and Avoidance Learning in Groups

Surprisingly, only a few studies have examined aversive learning in group settings, and of these, most have only used small social groups (i.e., dyads or triads) of rodent subjects to understand how learning about danger in groups affects individual behavior. Initially, these studies compared rats that experienced cue‐shock pairings by themselves versus rats that underwent fear conditioning in the presence of three cagemates that were separated by transparent dividers (Harb & Taylor, 2015; Fig. 1A). The transparent dividers allowed visual, auditory, and olfactory information to be transmitted while preventing physical interaction during shock delivery. When tested 24 hr later, the group‐conditioned rats showed significantly less freezing than the rats that underwent fear conditioning alone. This lower level of freezing persisted whether the rats were tested again alone or with their cagemates. Two follow‐up experiments in the same study revealed that removing olfactory cues (by using a zinc sulfate nasal wash before conditioning) or anesthetizing cagemates eliminated the effect of group conditioning.

Figure 1.

Figure 1

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Group fear‐conditioning paradigms. (A) Harb and Taylor (2015) conditioned rats either alone (top left) or in triads (bottom left) that were separated by transparent barriers. All were tested separately. Rats conditioned in triads showed significantly lower fear responses compared to rats conditioned alone. (B) Lee and Noh (2016) used dyads in stead of triads and also included a no‐shock comparison group. Rats conditioned in dyads showed significantly lower fear responses than rats conditioned alone. (C) Ferreira et al. (2019) conditioned rats alone or with a partner (left) and tested either alone or with a partner, or tested the dyad conditioned rats alone at test. Rats either conditioned or tested with a partner showed significantly less fear responses than rats conditioned alone. See Table 2 for more details on experimental design parameters. Red speaker represents tone conditioned stimulus; yellow lightning bolt represents shock unconditioned stimulus.

The finding that fear conditioning in a group setting reduces fear responses has been extended further using other approaches. For instance, a study in which fear conditioning occurred in freely interacting rat dyads reported significant reductions in freezing behavior and passive avoidance responses during subsequent memory tests (Lee & Noh, 2016; Fig. 1B). In another study that also used freely interacting animal dyads, rats underwent contextual fear conditioning either alone or in the presence of a cagemate, and then were tested individually or in the presence of the cagemate both 30 min and 24 hr later (Ferreira et al., 2019; Fig. 1C). Cagemate presence during the acquisition of contextual fear reduced freezing during the two memory time points, demonstrating that having a social partner present either during conditioning or during the memory test reduced fear memory. In the authors’ second experiment, rats were conditioned alone, then underwent a 10‐min extinction session either alone or with a cagemate, and then all rats were tested individually 24 hr later. Interestingly, although reduced freezing was observed during the extinction session in the presence of a cagemate, freezing levels were equivalent for both groups in the retention test, suggesting that a strong extinction memory was formed and retained despite the reduction in fear expression. Taken together, the above studies on fear learning in a group setting support the idea that the presence of a social conspecific reduces fear expression during and after aversive learning and that this effect requires group members to be conscious and able to use olfactory cues. Group fear learning also appears to exert a protective effect, similar to social coping, that is not simply a momentary reduction in fear but reflects a long‐lasting change.

One question that arises at this point is how stress might play a role during group‐based aversive learning. Studies show that rodents prefer to spend more time with a stressed familiar conspecific compared to unstressed unfamiliar conspecifics (Rogers‐Carter et al., 2018; Toyoshima et al., 2021), and a similar effect is observed during fear learning. After a rat has spent time with a recently fear‐conditioned conspecific, enhanced fear learning is observed (Knapska et al., 2010). A follow‐up study from the same research group assessed the effects of social presence during fear extinction (Gorkiewicz et al., 2023). After individual rats underwent fear conditioning, they underwent one session of fear extinction in the presence of a cagemate, followed by another session of fear extinction alone. Interestingly, these rats showed greater extinction of fear in the presence of the cagemate but not on the following day when undergoing extinction alone, implying that the social presence effect during the initial extinction session did not carry over to the next day. This suggests that having a social partner can reduce fear at any given moment, but it still may not facilitate memory consolidation of fear extinction. Together with the prior studies mentioned above, this finding may carry important implications for how social support during cognitive therapy for anxiety disorders may or may not improve the overall treatment outcomes.

Other studies have also examined the effects of social dimensions after fear learning, in which mice were fear conditioned individually and then tested the following day as freely interacting dyads in a different context (Ito et al., 2023; Fig. 2A). These studies also quantified the temporal synchronization of each dyad's freezing bouts, which can indicate how much of the freezing behavior might be due to the partner's freezing behavior, revealing much more information than just reporting the total time freezing of each mouse. To do this, the authors calculated how much more the two mice spent freezing simultaneously than would be expected by chance. Whereas both males and females showed significant freezing synchrony above chance, males synchronized more strongly than females. Further, removing visual cues eliminated freezing synchrony observed in most of the male, but not female, dyads, suggesting that males may rely more on visual information during social exchange of aversive learning. This finding aligns with recent work from my own laboratory using an observational platform‐mediated avoidance task (Obs‐PMA; see Fig. 3D), in which we found that male but not female rats tend to use more visual information to acquire threat avoidance responses when observing a highly experienced demonstrator that has already undergone multiple sessions of PMA (Ruble et al., 2026).

Figure 2.

Figure 2

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Other aversive learning paradigms using social dimensions. (A) Ito et al. (2023) conditioned mice alone (left) and then tested them in dyads. At test, mice showed synchronized freezing bouts with their partners. (B) Ruble et al. (2025) conditioned rats in the platform‐mediated avoidance (PMA) task in the presence of a non‐cagemate partner and then tested them separately. Freezing and avoidance were significantly higher in the absence than in the presence of their partner. See Table 2 for more details on experimental design parameters.

Figure 3.

Figure 3

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Paradigms using social transmission of fear. (A) Bruchey et al. (2010) created a Fear Conditioning by Proxy (FCbP) task and reported that some rats could learn fear from a cagemate that directly underwent auditory fear conditioning but resulted in variable levels of fear across rats at test. (B) Jeon et al. (2010) were the first to demonstrate observational learning of conditioned fear in which an observer mouse (black) witnessed a demonstrator mouse (grey) receive multiple footshocks through a transparent barrier. At test, the observer mouse showed increased fear responses. (C) Two subsequent studies (by Allsop et al., 2018, and Silverstein et al., 2024) implemented auditory fear conditioning and also showed mice or rats could successfully acquire conditioned fear at test. (D) Ruble et al. (2026) created an observational platform‐mediated avoidance task and revealed that Observer rats show the greatest avoidance at test after witnessing a novice rat performing the first day of PMA training. See Table 2 for more details on experimental design parameters.

A follow‐up study to the original social synchronization study above extended that work by examining how sex composition, prior stress, and partner familiarity modulate fear synchrony in dyads using the same procedure described above (Ito et al., 2025). After undergoing fear conditioning in individual animals, mice were tested in same‐sex or opposite‐sex dyads, with or without a brief restraint stress before testing, and with familiar or unfamiliar partners. In same‐sex male dyads, prior stress disrupted freezing synchrony. In contrast, same‐sex female dyads showed enhanced freezing synchrony when exposed to prior restraint stress. Partner unfamiliarity reduced synchrony in males but not females. Interestingly, opposite‐sex dyads showed high freezing synchrony that was resistant to both stress and unfamiliarity, suggesting that obtaining social information from the opposite sex may increase its saliency, heightening attention to the social information and thus making it more effective in understanding potential threats in the environment.

Similar findings have been reported in a modified version of the PMA task in which two rats undergo the task simultaneously (Kramer et al., 2025; Ruble et al., 2025; Fig. 2B). Just as in the standard PMA task in which rats avoid a tone‐signaled shock by stepping onto a platform across 10 days, pairs of non‐cagemate rats undergo the PMA task together. During this task, they have access to their own lever, food dish, and platform, while being separated by a perforated transparent barrier, allowing access to visual, auditory, and olfactory cues. When compared to those that learned the PMA task under solitary conditions, rats that underwent social partner PMA showed greater freezing responses during training. This was surprising, given that prior work has shown that the presence of another conspecific typically reduces freezing during fear conditioning (Ferreira et al., 2019; Harb & Taylor, 2015; Lee & Noh, 2016). However, the PMA task involves many more days of training (10 days, versus 1 day for fear conditioning) and greater task demands than fear conditioning. In addition, other studies have shown that during fear conditioning, when a rodent detects a social partner freezing or becoming suddenly immobile, this will also cause the rodent to freeze (Atsak et al., 2011; Cruz et al., 2020; Pereira et al., 2012; Pereira & Moita, 2016). This could explain why rats in the social partner PMA task show greater freezing in the presence of a social partner than rats in the solitary version of the PMA task.

We also asked whether the experience of the rat partner might influence avoidance learning by pairing rats that had already had 10 days of PMA training with rats with no prior PMA training (Ruble et al., 2025). Although we predicted that having an experienced partner might enhance PMA learning, there were no significant differences between rats with an experienced or a naive partner, suggesting that experience of a social partner alone was not effective enough to enhance avoidance learning. Moreover, we observed that compared to those undergoing solitary PMA training, rats that underwent social partner PMA training showed less food‐seeking behavior, evidenced by lower pressing during the intertrial interval. This suggested that rats were attending less to the food, likely because they were attending more to their social partner. Future studies are currently trying to answer this question by determining whether any behavioral synchrony between rat partners contributes to learning during the PMA task.

Social Transmission of Fear and Avoidance: Fear Conditioning by Proxy and Observational Learning

Fear conditioning by proxy is an aversive learning paradigm in which individuals learn about danger through social transmission of information but without directly observing an individual undergoing an aversive experience (Bruchey et al., 2010; Fig. 3A). For this procedure, the demonstrator has already undergone fear conditioning away from other conspecifics before the social interaction begins. The observer then interacts freely with the conditioned demonstrator and is later exposed to the conditioned stimulus (CS) in the absence of the shock unconditioned stimulus (US). Any fear response the observer acquires is thought to come from the social cues the observer gained during the interaction with the previously conditioned demonstrator. The resulting fear learning in this paradigm is more variable than in observational fear conditioning: some observers show no CS‐induced freezing, whereas other observers show high levels of CS‐induced freezing (Bruchey et al., 2010; Monfils & Agee, 2019). Interestingly, the best predictor of whether an observer acquires this fear response is the amount of pro‐social contact it makes with the demonstrator during the session (Bruchey et al., 2010). When dominance status was taken into consideration, subordinate rats reliably acquired fear from dominant demonstrators, but not vice versa (Jones & Monfils, 2016).

Another learning paradigm that evaluates the social transmission of information is observational fear learning, which relies on how individuals acquire new behaviors by observing how other conspecifics learn and react to meaningful stimuli in their environment. This is critical for safety and survival because during situations involving threat, harm, or punishment, this learning process allows individuals to acquire the meaning of stimuli that predict these aversive outcomes without ever being exposed to them directly and to subsequently use that information to safely navigate their environment. This type of learning has been studied for decades in rodents (Keum & Shin, 2019; Kitamura et al., 2024; Monfils & Agee, 2019), nonhuman primates (Putnam & Chang, 2021; for cross‐species review, see Burgos‐Robles et al., 2019), and humans (Debiec & Olsson, 2017; Skversky‐Blocq et al., 2021). More recently, researchers have begun to reveal the neural mechanisms that guide observational aversive learning, specifically for fear conditioning. In the basic procedure, an observer and demonstrator are placed in adjacent compartments separated by a transparent barrier, which allows transmission of visual, auditory, and olfactory cues between the two animals. For observational conditioning, the demonstrator receives a series of footshocks while the observer watches. In some designs, auditory cues are also presented with the footshocks (Allsop et al., 2018; Silverstein et al., 2024; see Fig. 3C). When the observer is later tested alone, either in the same context or in response to the auditory CS, the observer shows fear responses that it never learned through direct experience.

The initial studies on this paradigm used context fear conditioning as the basis of learning, in which an observer witness a demonstrator experiencing a series of footshocks within a chamber (Jeon et al., 2010; Jeon & Shin, 2011; Fig. 3B). Observers were tested in the same context 24 hr later and displayed freezing to the context. This was the first demonstration that mice could form a contextual fear memory without direct experience of the shock US. The authors also reported that familiar conspecifics were more effective at transmitting fear through observation. Additional experiments further showed that replacing the transparent barrier with an opaque one reduced but did not eliminate observational learning, suggesting that visual cues are not the only source of sensory information that is used to acquire observational fear.

Later studies employed observational learning of auditory fear conditioning (Allsop et al., 2018; Silverstein et al., 2024) in which the observer witnessed the demonstrator receive tone‐signaled footshocks, enabling the fear response (also freezing in this case) to be time‐locked to a discrete cue instead of the context (Fig. 3C). This design is advantageous because observers formed a fear memory that could be tested in a novel context, which would rule out contextual generalization as an explanation of the observed fear. Both contextual and cued observational fear tasks do hold some ethological value. In the wild, an animal that detects another animal's distress, through vocalizations, pheromones, or visible defensive behavior, can gain information about danger that it has not directly encountered. This is a safer way to learn about danger and a common way that information spreads through social groups. Likewise, humans regularly acquire fear and anxiety through observation and social transmission, a process that can contribute to the development of post‐traumatic stress disorder (PTSD) and other anxiety disorders some individuals (Debiec & Olsson, 2017; Delgado et al., 2006).

Other studies have assessed observational aversive learning during active avoidance using hurdle‐jumping or shuttle‐avoidance tasks. Early research reported that rats (del Russo, 1975), cats (John et al., 1968), and nonhuman primates (Presley & Riopelle, 1959) perform active avoidance better if learned through observation, compared to direct learning of active avoidance. A recent study modified the platform‐mediated avoidance (PMA) task to study observational avoidance learning, in a protocol in which an observer rat witnessed a demonstrator rat undergo Day 1 (a novice demonstrator) or Day 10 (an experienced demonstrator) of PMA training (Ruble et al., 2026; Fig. 3D). During the PMA task, rats learn to avoid a tone‐signaled footshock by stepping onto a safe platform, which comes at a cost of lever pressing for a sucrose reward. Observers that witnessed novice demonstrators showed greater avoidance learning than observers that witnessed experienced demonstrators, suggesting that novice rather than experienced demonstrators offer more information to observers about the PMA task. Interestingly, this effect was only observed in rats that underwent shock pre‐exposure before observation, which may have enhanced observers’ learning about the tone‐shock association, something that subsequently enabled learning about the adaptive behavior of avoiding shock by stepping onto the platform.

During the Observational PMA task, a subset of Observer rats that witnessed a Novice Demonstrator showed greater avoidance learning at test compared to rats that observed an Experienced Demonstrator (Ruble et al., 2026). We interpreted this finding as rats observing novice rats experiencing all aspects of the task for the first time are likely to show a larger variety of behaviors, such as freezing, reacting to the shock, etc. This is similar to our finding during the social partner PMA task that having either an experienced partner or a novice partner led to similar levels of learning during training.

Some research on observational fear conditioning argues that shock pre‐exposure, typically a single footshock given before the observation session, is necessary for the observer to understand the ongoing experience of the demonstrator; otherwise, observational learning may not occur (Allsop et al., 2018; Atsak et al., 2011; Kim et al., 2010; Sanders et al., 2013). Specifically, the claim is that the observer needs an experiential reference point to associate the demonstrator's distress with the dangerous outcome. Other research argues that any prior shock exposure introduces direct learning that interferes with the observational learning, making it impossible to isolate the effect that the socially observed information has on learning (Chen et al., 2009; Ito et al., 2015; Ito & Morozov, 2019; Jeon et al., 2010; Ramesh et al., 2025; Silverstein et al., 2024; Yusufishaq & Rosenkranz, 2013). Both sides of the argument have empirical support: observers show stronger and more reliable fear learning when they have had prior shock exposure (Allsop et al., 2018), but observational aversive learning tasks without any shock pre‐exposure have also demonstrated observational learning of conditioned fear (Jeon et al., 2010; Silverstein et al., 2024) or active avoidance (Ruble et al., 2026), albeit to a lesser extent. Other studies have either used greater shock intensity or administered a larger number of shock trials to the demonstrator in order to increase observational freezing (for review, see Burgos‐Robles et al., 2019).

A potential solution for this problem could be the use of non‐shock aversive stimuli as the pre‐exposure experience, for example exposure to a predator odor or mild stressors in the homecage (e.g., wet bedding, loud background noise). These stimuli would elicit an aversive experience in the observer without introducing direct shock learning and could also eliminate the possibility that shock exposure alone is driving the effects of learning. Moreover, if using non‐shock aversive stimuli was compared with using a shock pre‐exposure before observational learning, this could confirm whether shock‐pre‐exposed rats use direct learning of shock to drive observational learning. However, this may depend on the type of aversive stimuli used as the pre‐exposure experience. For example, a study using a swim stress test 24 hr before observation fear learning showed it did not enhance fear in observers, suggesting that it is the shared experience (not just any aversive stimulus or experience that increases stress) that enhances learning (Sanders et al., 2013). Future work can follow up on this question using other aversive experiences or stimuli.

Taken together, these paradigms demonstrate that fear in response to specific cues and contexts can be acquired through the transmission of social information, either through observation or through subsequent social interactions. Learning about danger without directly experiencing danger is a powerful tool that social animals can use to learn about the dangers of the world in a safe way. Key variables such as the source of social information and prior experience can greatly affect the strength of the outcome of what is learned through social information, which will be discussed in the next section on relevant social factors influencing aversive learning.

Social Fear Conditioning

Another social dimension of interest for aversive learning is when the social interaction itself becomes aversive. This type of learning has been termed social fear conditioning (SFC) or social threat conditioning (STC). During SFC/STC, mice receive footshocks during active social investigation of an unfamiliar conspecific (Toth et al., 2012; Fig. 4A). After conditioning, mice developed increased freezing and avoidance of the social stimulus that was paired with shock. Toth and colleagues (2012) also showed that first‐line pharmacological treatments used for treating social anxiety disorder in humans (both diazepam and paroxetine) reduced social fear in this paradigm, making it a clinically relevant model for the study of social anxiety disorder.

Figure 4.

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Social fear conditioning. (A) Toth et al. (2012) paired shock with approaching a conspecific mouse. At test, mice showed increased freezing and avoidance of the stimulus mouse. (B) Lozano‐Ortiz et al. (2025) used this task to investigate differential SFC and found that mice could discriminate between a mouse paired with shock and another mouse unpaired with shock. But subsequent testing showed the test mouse (black) began generalizing across all mice and showed social avoidance. See Table 2 for more details on experimental design parameters.

The SFC paradigm was subsequently enhanced in several ways (Lozano‐Ortiz et al., 2025; Velazquez‐Hernandez et al., 2024; Xu et al., 2019; Zheng et al., 2021). First, the SFC procedure was automated to reduce behavioral variability by reliable trial‐by‐trial detection of social contacts between mice and by delivering footshocks with precise timing. Second, to rule out any potential fear due to contextual features or other cues in the environment (e.g., the cage of the stimulus mouse), the conditioning chamber was modified by placing a second cage that was identical to the cage containing the stimulus mouse but that remained empty (Lozano‐Ortiz et al., 2025; Velazquez‐Hernandez et al., 2024; Xu et al., 2019; Zheng et al., 2021). Third, the duration of the conditioning session was increased because prior work has indicated that remaining near a social stressor enhances memory of the aversive experience (Golden et al., 2011).

Unsupervised machine learning computational approaches were used to reveal novel behavioral sequences the conditioned mice were expressing during SFC (Velazquez‐Hernandez et al., 2024). The authors found unique behavioral sequences of the body and tail movements across all mice, and significantly different behavioral sequences between females and males. Another study using an automated version of SFC also found increased darting and stretching postures following SFC (Lozano‐Ortiz et al., 2025). Darting was reported to be more prevalent in female than in male mice, similar to other studies that have reported more darting in female rats during auditory fear conditioning (Gruene et al., 2015). Interestingly, when shock was paired with only one of two mice used as social stimuli, the conditioned mice successfully discriminated the mouse paired with shock and mostly avoided that mouse stimulus during the training session, but after another sociability test, the conditioned mice exhibited generalized avoidance of both social stimuli (Lozano‐Ortiz et al., 2025; Fig. 4B).

Key Considerations for Social Dimensions in Aversive Learning

Collectively, the studies described above highlight many key variables that should be considered when utilizing social dimensions within an aversive learning paradigm. These variables can affect how an individual learns about danger through social information, including (but not limited to) relevant prior experience, the relationship between the individual learning about danger and the conspecific(s) from which they are learning, and specific features or internal states of the conspecific. Table 1 describes these variables and their implications for aversive learning.

Table 1.

Key Considerations when Implementing a Social Dimension During Aversive Learning Paradigms a

Key considerations for social dimensions in aversive learning

1: What is the influence of animals’ prior experiences?

If one or more of the conspecifics has prior experience with the task or some aspect of the task (e.g., shock exposure), this experience may enhance learning.

2: What is the relationship between the animals?

Cagemates, siblings, mating partners or animals with some other bond can influence learning.

3: What features/traits among the animals may affect learning?

Age, strain, sex, and dominance status, and other fixed traits, might affect learning.

Transient features may also influence learning, such as emotional, physiological state (e.g., hormones, stress, hunger).

4: Are there novel behaviors occurring that are not typically observed during individual aversive learning?

New behaviors (e.g., huddling if freely interacting, or mimicking if divided by transparent barrier) or behavioral sequences might emerge when adding a social dimension which might influence learning.
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a

Several questions and potential outcomes are listed that should be considered when designing experiments.

WHERE DO WE GO FROM HERE?

The studies reviewed here show that a variety of social dimensions can shape fear learning and behavior. Moving forward, the field needs to develop innovative strategies to further incorporate social elements into animal models to gain a better understanding about fear learning in more naturalistic ways. Several promising directions are already emerging that are discussed below.

Standard laboratory paradigms that utilize small chambers and artificial cues to test isolated animals are useful for reducing variability across laboratory‐based studies. Yet the studies described thus far have used a wide selection of behavioral designs that vary in apparatuses, number of trials presented, shock parameters, behavioral measures, etc. (see Table 2), which make it difficult to draw strong conclusions about each of their findings across the literature. Furthermore, these designs strip away much of the complexity in which fear and coping operate in realistic situations. Larger, more complex environments that allow social interaction, exploration, and escape give animals the opportunity to express a fuller range of behavioral strategies, including social ones. Tasks that incorporate predator‐relevant stimuli, such as the approach‐avoidance task using a “robogator” stimulus (Choi & Kim, 2010) or predator odor (Fernandez‐Leon et al., 2021; Rodriguez‐Romaguera et al., 2020) are of particular interest because they engage defensive behavioral systems that may be more ecologically representative. Conflict tasks such as the Risk‐Reward Interaction task (Kyriazi et al., 2018), the Crossing‐Mediated conflict task (Illescas‐Huerta et al., 2021), and the high‐conflict PMA task (Bravo‐Rivera et al., 2021; Ramirez et al., 2024) that place animals in motivational conflict between threat and reward can be extended to ask how the presence of a conspecific shifts the balance. Finally, to understand how social partners might influence safety learning and dynamic escape behavior, social dimensions could be incorporated into recently developed paradigms, including the fear‐safety‐reward discrimination task (Greiner et al., 2019; Sangha et al., 2013) and escape from threat in complex environments (Wang et al., 2021).

Table 2.

Key Studies Implementing Social Dimensions in an Aversive Learning Task with Animal Information, Conditioned and Unconditioned Stimulus Parameters, and Number of Trials Indicated

Study Animals CS parameters Shock US parameters Design
Species Strain Sex Type Duration Intensity Duration No. trials
Harb & Taylor (2015): Auditory FC Rat Sprague Dawley Male only Tone 30 sec 0.8 mA 2 sec 1 or 3
Lee & Noh (2016): Auditory FC Rat Sprague Dawley Male only N/A (context) 1.0 mA 5 sec 1
Ferreira et al. (2019): Auditory FC Rat Wistar Male only N/A (context) 0.5 mA 2 sec 3
Ito et al. (2023): Auditory FC Mouse C57BL/6N and 129SvEv / C57BL/6N Male and female Tone 30 sec 0.5 mA 0.5 sec 4
Ruble et al. (2025): SP‐PMA Rat Sprague Dawley Male and female Tone 30 sec 0.4 mA 2 sec 9 (×10 days)
Bruchey et al. (2010): FCbP Rat Sprague Dawley Male only Tone 20 sec 0.7 mA 0.5 sec 3
Jeon et al. (2010): OFL Mouse C57BL/6J Male only N/A (context) 1.0 mA 2 sec 25
Allsop et al. (2018): AV OFL Mouse C57BL/6J Male only Tone and light 20 sec 1.0‐1.5 mA 2 sec 30
Silverstein et al. (2024): Auditory OFL Mouse C57BL/6J Male only Tone 30 sec 1.0 mA 2 sec 30
Ruble et al. (2026): Obs PMA Rat Sprague Dawley Male and female Tone 30 sec 0.4 mA 2 sec 9
Toth et al. (2012): SFC Mouse CD1 Male only N/A (US paired with stimulus mouse) 0.7 mA 1 sec 2‐5
Lozano‐Ortiz et al. (2025): SFC Mouse C57BL/6J Male and female N/A (US paired with stimulus mouse) 0.4 mA 0.5 sec Varied during 20‐min session
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AV, audiovisual; CS, conditioned stimulus; FC, fear conditioning; FCbP, fear conditioning by proxy; N/A, not applicable; Obs PMA, observational platform‐mediated avoidance; OFL, observational fear learning; SFC, social fear conditioning; SP‐PMA; social partner platform‐mediated avoidance; US, unconditioned stimulus.

Such future directions will also require methodological advances in behavioral analysis. Tracking multiple animals simultaneously requires sophisticated tools to measure dynamic behavioral interactions in real time and to quantify the moment‐to‐moment influence of one animal's behavior on its partner's. Advances in multi‐animal pose estimation and automated behavioral classification are making this increasingly feasible (Chen et al., 2023; Goodwin et al., 2024; Mathis et al., 2018; McTaggart et al., 2024; Weinreb et al., 2024); therefore, integrating these tools into social aversive learning paradigms is an important next step.

Social transmission and social coping strategies are other directions that are highly relevant to better understanding of how incorporating social elements could lead to more effective neuropsychiatric treatments. Preclinical studies might investigate how a resilient individual could socially buffer a susceptible individual and, if it does, through what behavioral and neural mechanisms. These lines of work move beyond asking how social context modulates aversive learning to asking how social interaction shapes the capacity to cope. This is a distinction with significant implications for understanding behavioral resilience and for developing socially based interventions for anxiety disorders in humans. For example, human studies have shown that a strong social support system can reduce anxiety (Boscarino, 1995; Brewin et al., 2000; Platt et al., 2014), but future studies should consider other social factors that may contribute to effective behavioral treatments for anxiety and other neuropsychiatric disorders.

Finally, many of the studies described here and others have begun to elucidate the neural mechanisms that guide aversive learning under these various social conditions. For example, Leblanc and Ramirez (2020) reviewed some recent work on social learning in rodents, focusing on the need to uncover a social memory engram. Moreover, many studies now support the idea that many of the major hubs that are central for aversive learning under solitary conditions are also recruited under social conditions (Olsson et al., 2020). However, future work should continue investigating the neural mechanisms of the social regulation of fear to further define these circuits and how they may differ among these various aversive learning paradigms implementing social dimensions.

AUTHOR CONTRIBUTIONS

Maria M. Diehl: Conceptualization; investigation; funding acquisition; writing—original draft; writing—review and editing.

CONFLICT OF INTEREST

The author does not declare any conflicts of interest.

ACKNOWLEDGMENTS

The author thanks Drs. Anthony Burgos‐Robles and Shana Silverstein for helpful comments on the manuscript.

Diehl, M. M. (2026). Beyond fear learning in isolation: Integration of social dimensions into aversive learning. Current Protocols, 6, e70424. doi: 10.1002/cpz1.70424

Published in the Neuroscience section

DATA AVAILABILITY STATEMENT

Data sharing not applicable to this article as no datasets were generated or analyzed during the current study.

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Data sharing not applicable to this article as no datasets were generated or analyzed during the current study.

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