Alcohol & Alcoholism Vol. 40, No. 5, pp. 373378, 2005 doi:10.1093/alcalc/agh177Advance Access publication 4 July 2005

CAN ALCOHOL LEAD TO INHIBITION OR DISINHIBITION? APPLYING ALCOHOLMYOPIA TO ANIMAL EXPERIMENTATION

NAOMI K. GRANT* and TARA K. MACDONALDDepartment of Psychology, Queens University Kingston, Ontario, K7L 3N6, Canada

(Received 24 June 2004; first review notified 22 February 2005; in revised form 30 May 2005; accepted 31 May 2005; advance access publication 4 July 2005)

Abstract Aims: Animal experimentation often demonstrates that alcohol leads to disinhibited behaviour, such as increasedaggression, increased social behaviour, or increased impulsivity. However, human experimentation demonstrates that alcohol can haveeither disinhibiting or inhibiting effects on behaviour, depending on salient environmental cues. Our aim was to illustrate how alcoholmyopia theory could be applied to the literature assessing the effects of alcohol on behaviour in animals. Methods: The effects ofalcohol on animal behaviour were reviewed in several domains, including aggression, social behaviours, and impulsivity. Suggestionsfor testing alcohol myopia with animal research paradigms were provided. Results: Current animal research paradigms are oftendesigned in such a way that alcohol myopia cannot be tested. To test alcohol myopia, we recommend manipulating the salience of bothimpelling and inhibiting environmental cues. Conclusions: Disinhibition alone cannot explain alcohols effects on behaviour. Wecontend that alcohol myopia theory helps to explain some contradictory findings in the human and animal literature. We encourageanimal researchers to develop research paradigms to provide tests of alcohol myopia.

The Author 2005. Published by Oxford University Press on behalf of the Medical Council on Alcohol. All rights reserved

*Author to whom correspondence should be addressed at: Department ofPsychology, Queens University Kingston, Ontario, K7L 3N6, Canada. Tel.:+1 613 533 2873; Fax: +1 613 533 2499; E-mail: 9nkg@qlink.queensu.ca ortmacdon@post.queensu.ca

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INTRODUCTION

How does alcohol affect behaviour? One popular notion is thatalcohol causes disinhibition, or a release of natural impulses byeliminating learned inhibitions (Critchlow, 1986). For example,alcohol is used as a social lubricant at parties, which is evidenceof this strong belief in the disinhibiting effects of alcohol onpositive social behaviours. However, alcohol is also believed toincrease antisocial impulses, such as aggression or risk-taking.

Disinhibition alone, however, cannot account for the complex,and sometimes contradictory research findings. On the one hand,animal research often demonstrates that alcohol leads todisinhibited behaviour, such as increased aggression (Miczeket al., 1993), impulsivity (Poulos et al., 1998; Evenden and Ryan,1999), and playfulness (Varlinskaya et al., 2001), consistent withthe idea that alcohol is a general disinhibitor. The humanliterature has also been guided to a large extent by disinhibitiontheory. However, recent research shows that alcohol does notalways lead to disinhibited behaviour, but can either inhibit ordisinhibit behaviour, depending on environmental cues.

ALCOHOL MYOPIA THEORY

Why does alcohol cause disinhibited behaviour in some casesbut not in others? An explanation is offered by alcohol myopia,which states that alcohol limits cognitive capacity such thatintoxicated individuals tend to focus on cues in the environmentthat are most salient (Steele and Josephs, 1990). Alcoholmyopia postulates that intoxicated individuals are unable toattend to all relevant cues simultaneously because of thelimitation of cognitive capacity associated with alcoholintoxication. In other words, alcohol produces a myopic effectcausing individuals to attend primarily to, and hence be moreinfluenced by, salient environmental cues at the expense of less

salient cues. Individuals who are not intoxicated, however, arenot as easily influenced by salient cues because they are betterable to attend to all the relevant information in the environment.

Evidence to support alcohol myopia has been found in thedomain of health-relevant behaviours, such as drinking anddriving or engaging in unprotected sex. MacDonald et al.(1995) found that when asked about their attitudes towardsdrinking and driving, both sober and intoxicated individualsreported very negative attitudes. However, when the wordingof the question included an impelling cue to drink and drive(e.g. having to drive only a short distance), intoxicatedindividuals reported less negative attitudes than soberindividuals. This pattern is consistent with alcohol myopiabecause intoxicated participants were influenced by theimpelling cue, but sober participants were not.

Intentions towards engaging in unprotected sex have alsobeen shown to be influenced by salient inhibiting cues suchthat intoxicated individuals actually reported more prudentintentions compared with sober controls (MacDonald et al.,2000). It is important to note that disinhibition theory cannotaccount for these findings; it cannot explain situations in whichintoxicated individuals behave more prudently than soberindividuals. If alcohol is a general disinhibitor, intoxicatedindividuals should always exhibit disinhibited behaviour.

Findings emerging from the human literature present adifferent picture than those in the animal literature. Althoughconventional wisdom might argue that alcohol will alwayslead to disinhibited behaviour, it is clear that this explanationis less than satisfactory. Indeed, researchers are aware that therelationship between alcohol and behaviour is complex, andcannot be accounted for by simple disinhibition (Miczek et al.,1993). We argue that alcohol myopia might resolve some ofthe complexities and contradictory findings.

APPLYING ALCOHOL MYOPIA TO THEANIMAL LITERATURE

Animal research has virtually ignored alcohol myopia. Indeed,most research paradigms in the animal literature do not allow

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for a true test of alcohol myopia. To test alcohol myopia,both impelling and inhibiting cues should be present. Ideally,the salience of these cues is also manipulated. In manyanimal research paradigms, the salient environmental cuesare generally impelling ones, i.e. ones that would elicitdisinhibition. For example, in testing the influence of alcoholon aggression, some researchers use a shock chambersituation (Tramill et al., 1981; Davis et al., 1993). The animalis restrained in a chamber and aggression is elicited byshocking the animal. The shock can be seen as an impellingcue to act aggressively. Therefore, when an alcohol-treatedanimal behaves more aggressively, is it because of thedisinhibiting effects of alcohol, or is it because the shock isa salient, impelling cue to act aggressively? To test bothdisinhibition theory and alcohol myopia, two cue conditionswould need to be present in the study. For instance, one couldcompare conditions in which shock is either present or absent.If aggression increases regardless of the presence of shock,this would suggest that alcohol is simply producingdisinhibition. If, however, the increase in aggression inalcohol-treated animals occurs only when the shock is present,that would suggest that the shock is an impelling cue to behaveaggressively.

Cutler et al. (1975) examined the effects of alcohol onsocial behaviours in mice and found that alcohol onlyincreased behaviours that were already stimulated by thetest situation. When the mice were subjected to a new envi-ronment, alcohol increased exploratory behaviour. However,alcohol-treated mice which were not subjected to the newenvironment showed no differences in their exploratorybehaviour from untreated mice. Similarly, in a territorialsituation, when territories had not been established, alcohol-treated mice became dominant over the entire enclosure.When territorial boundaries were established, however,alcohol-treated mice were no more likely to dominate theenclosure than the non-treated mice. Although the authorsview these effects in terms of disinhibition, we contend that abetter explanation might be alcohol myopia. When cuesdesigned to elicit a particular behaviour were present, alcohol-treated mice showed increases in that behaviour. When thesecues were not present, alcohol-treated mice did not differfrom the non-treated mice, which is exactly what alcoholmyopia predicts.

Although the results of the Cutler et al. (1975) study areconsistent with alcohol myopia, only impelling cues wereincluded in the study. It is important to note that disinhibitiontheory and alcohol myopia make opposing predictions wheninhibiting cues are present. Disinhibition theory predicts thatbehaviours that would normally be inhibited, such asaggression or impulsivity, will increase under the influence ofalcohol. Alcohol myopia predicts that, if powerful inhibitingcues are present, alcohol could cause animals to behave evenless aggressively or impulsively than they would otherwise.The finding that alcohol intoxication can lead to moreinhibited behaviour has important implications for research,especially for studies on humans, as many of the behavioursassociated with alcohol are potentially harmful (e.g. drinkingand driving, risky sexual behaviour, and aggression).Furthermore, including tests of alcohol myopia in animalresearch will lead to a better understanding of how alcoholaffects behaviour.

The purpose of this paper is to explain how alcohol myopiacould be tested within animal research paradigms. First, wewill review the animal literature and main findings for thebehavioural domains of aggression, social behaviour, andimpulsivity. We believe it would be useful to considerapplying alcohol myopia to these particular domains for tworeasons. First, the effects of alcohol on behaviour have beenwidely studied in these areas. Second, the behaviours in thesedomains are relatively complex, making them more pertinentto the human alcohol myopia literature on social decision-making. We will suggest ways to test alcohol myopia theorywithin each of these domains, and finally, we will explain thespecific predictions that follow from alcohol myopia andcompare these predictions with those from a disinhibitionperspective. We will primarily discuss the findings regardingthe short-term effects of alcohol at low and moderate doses,because these are most comparable with the studies conductedon humans. Furthermore, although a biphasic doseeffectcurve of alcohol on behaviour has been shown such that lowerdoses often cause increases and higher doses cause decreasesin a particular behaviour (Miczek et al., 1993), it is possiblethat the effects at high doses might be owing to the sedativeeffects of alcohol, which is less pertinent to an alcohol myopiaperspective.

AGGRESSION

The effects of alcohol on aggression have been studiedextensively. To date, studies involve placing the animal in asituation where aggressive behaviour would normally beelicited (e.g. a residentintruder paradigm) and observing theeffects of alcohol. For example, when resident ciclid fishconfronted intruder fish, moderate doses of alcohol producedincreases in attacks (Peeke et al., 1973). Similarly, in responseto their mirror image, Siamese fighting fish treated with lowdoses of alcohol increased their aggressive displays (Rayneset al., 1968; Raynes and Ryback, 1970).

Using residentintruder paradigms, low doses of alcoholhave been found to increase aggressive behaviour in mice(Krsiak, 1976), increase or decrease aggression depending onwhether confrontations took place in neutral or home cages,(Miczek and ODonnell, 1980), and to have no effects onaggression (Lagerspetz and Ekqvist, 1978). It is not surprisingthat alcohol often causes increases in aggression in theseparadigms. The finding that alcohol-treated animals behavemore aggressively when they are confronted with an intruderis consistent with alcohol myopia because the intruder actsas an impelling cue to behave aggressively. Alcohol myopiawould predict that as animals become intoxicated, they will beunable to attend to all the relevant cues and will focus on themost salient cues, which in these studies would be the intruder.However, it is not clear whether disinhibition or alcoholmyopia is the process by which the alcohol-treated animalsbehaved more aggressively because disinhibition theorywould predict a similar increase in aggression. Are theseeffects the result of the disinhibiting properties of alcohol? Ordoes the intruder act as an impelling cue to act aggressively,which becomes the most salient cue for the alcohol-treatedanimals, thereby eliciting aggressive behaviour? It is difficultto test alcohol myopia within this paradigm because an

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intruder generally leads to aggressive behaviour by theresident. Even comparing the residentintruder confrontationstaking place in neutral versus home cages is difficult becausethe rate of aggression is often so high in the home cage that aceiling effect might occur.

In the residentintruder paradigm, resident rats typicallydisplay a specific pattern of aggressive behaviour resulting indefeat of the intruder, who eventually displays submissivebehaviour. Although alcohol myopia is difficult to test in thisparadigm, some indirect support for alcohol myopia can befound in a study by van Erp and Miczek (1997). Resident ratsthat showed large increases in aggression after self-administration of alcohol were classified as high in alcohol-heightened aggression. The authors speculate that alcoholmade these rats unresponsive to signals of submission by theintruder rat. These signals of submission by the intruderusually lead to decreased attacks by the resident. The idea thatthe alcohol-treated rats attended to the salient cue to actaggressively (an unfamiliar intruder) at the expense of lesssalient inhibiting cues (subtle signs of submission) is consistentwith alcohol myopia because it suggests that these rats werenot able to attend to all the relevant cues simultaneously.

Some researchers place the animal in a competitivesituation and observe the effects of alcohol on aggression. Inthese studies, animals are categorized based on their socialstatus, which appears to interact with alcohol. Low andmoderate doses of alcohol have been shown to increaseaggression in dominant rats (Miczek and Barry, 1977) and indominant squirrel monkeys (Winslow and Miczek, 1985).However, Pettijohn (1979) found that during a competitionbetween three dogs over a bone, low doses of alcoholincreased aggression in subordinate dogs but reducedaggression in higher ranking dogs. If alcohol were simplydisinhibiting aggressive behaviour, why would alcoholproduce different effects depending on social status? It ispossible that environmental cues function differently foranimals of different ranks. Perhaps alcohol leads subordinatedogs to attend to the salient bone at the expense of the lesssalient social hierarchy. If disinhibition theory were true, allalcohol-treated animals should respond the same way whenprovoked. Instead, the meaning of the cues may varydepending on the social status of the animals; so animals willrespond differently in a competitive situation depending ontheir social status, which is consistent with alcohol myopia.

Shock or pain has also been used to elicit aggression inalcohol-treated animals. For example, Weitz (1974) found thatalcohol increased fighting behaviour in pairs of male rats whenelectric foot shock was employed. In contrast, Tramill et al.(1980) found that low doses of alcohol decreased aggressiontowards a lever when single-restrained rats were shocked. It isdifficult to compare these two studies because the cues aredramatically different. While shock was employed in bothstudies, and may have acted as a cue to behave aggressively, thetarget of aggression was another male rat in the study of Weitz(1974) and a lever in the study by Tramill et al. (1980). It isquite possible that another male rat could provoke the alcohol-treated rat such that it becomes an additional cue to actaggressively. It is hard to imagine, however, how a lever in thisinstance could act as an impelling or an inhibiting cue. To testalcohol myopia, it would be important to have two types of cueconditions in the same study.

To sum up, alcohol studies on several types of animals havegenerally shown that low and moderate doses of alcohol causeincreases in aggression. However, most of this research involvessubjecting the animals to situations where impelling cues to actaggressively are present, such as residentintruder confronta-tions, competitive situations, or shock chambers. To test alcoholmyopia, it is important to include inhibiting cues, or ones thatwould encourage an animal to behave non-aggressively.

SOCIAL BEHAVIOURS

The effects of alcohol on social behaviours have also beenstudied in animals. Alcohol has been shown to increase playbehaviours in monkeys (Cressman and Caddell, 1971; Crowleyet al., 1974). In contrast, Krsiak and Borgesova (1973) foundthat alcohol decreased all social activities in rats. This researchdoes not provide a test of alcohol myopia because theenvironmental cues were not measured or manipulated. Forexample, the studies on monkeys involved placing the alcohol-treated monkey with three other monkeys. The resultinginteractions were then observed. It is impossible to control thebehaviour of the other monkeys making it difficult to knowwhether the salient cues available to the alcohol-treatedmonkey were impelling cues to act sociably, impelling cues toact aggressively, or a combination of the two.

Varlinskaya et al. (2001) also investigated the effects ofalcohol on social behaviours in rats. Alcohol-treated rats wereeither exposed to social stimuli (littermate of same gender)or non-social stimuli (a cotton ball) placed in the testingchamber. They found that low and moderate doses of alcoholincreased social behaviours in the social condition (animpelling cue to act sociably), but failed to do so in the non-social condition (no impelling cue), which is consistent withalcohol myopia. A better test of alcohol myopia would includean inhibiting cue, or a cue to act non-sociably. For example,one could compare the responses of alcohol-treated rats to alittermate with their responses to an unfamiliar rat. Accordingto alcohol myopia, alcohol should increase prosocialbehaviours in the presence of a littermate (an impelling cue),but should decrease prosocial behaviours in the presence of anunfamiliar rat (an inhibiting cue). However, in this paradigm,these cues might be confounded with other aggression cues,such as dominant and subordinate positions.

Studies of the effects of alcohol on social behaviours havegenerally shown that alcohol tends to increase socialbehaviours. The challenge for testing alcohol myopia in thisdomain is that cues are difficult to measure or manipulate.Although it makes sense to test social behaviours in responseto another animal or a group of animals, it is hard to controlthe behaviour of the other animals. To test alcohol myopia,one would need to design a study in which cues could bemanipulated in some way to produce impelling cues (or cuesto behave prosocially) in one condition and relativelyinhibiting cues (or cues to behave antisocially) in another.

IMPULSIVITY

To test alcohols effects on impulsivity, animals are oftenpresented with a choice between a small, immediate reward or

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a large, delayed reward. The index of impulsivity, then, isbased on the number of times the animal chooses theimmediate reward. Poulos et al. (1998) trained rats to respondto a T-maze in which one arm led to 2 food pellets and theother arm led to 12 food pellets. Delay was then introduced forthe large reward. They found that alcohol dose-dependentlyincreased impulsivity, or choice of the arm that led to thesmall, immediate reward. Tomie et al. (1997) had rats choosebetween two levers, one producing a small, immediate rewardand the other producing a large, delayed reward. They foundthat low and moderate doses of alcohol increased thepreference for the small, immediate reward (see also Evendenand Ryan, 1999).

Although these findings are consistent with disinhibition,they are also consistent with alcohol myopia. Foodadministered with no delay is probably more salient thandelayed food, and alcohol myopia would predict that alcohol-treated animals should focus on the salient cue (the immediatereinforcer) at the expense of the less salient cue (the delayedreinforcer). It would be useful to design a study in which thelarge, delayed reward was made more salient. For example,one might modify the delay-of-reward paradigm to have thelarge reward inaccessible to the animal during the delay butstill in full view, and have the small reward hidden from view.Alcohol myopia would predict that the alcohol-treated animalsshould show a preference for the large, delayed reward at theexpense of the small, immediate reward when the large rewardis a more salient cue. However, if disinhibition theory is true,animals should be unable to delay gratification, and shouldimpulsively choose the small immediate reward in favour ofthe large, delayed reward.

One T-maze study of alcohols effects on impulsivity didprovide a test of alcohol myopia. P. C. Darling, T. L. Pinder,K. G. C. Hellemans, T. A. Paine and M. C. Olmstead (2003)trained rats in a T-maze in which one arm led to a small,immediate reward and one arm led to a large, delayed reward(Unpublished data). In the first study, alcohol increasedimpulsivity as measured by choice of the immediate reward.In the second study, a light predicted either the smallimmediate or the large delayed reward, but the arm paired withthe light varied across trials. In this study, alcohol caused anincrease in the tendency of the rats to run towards the lit arm,regardless of whether it was associated with a small or largereward. Control rats did not attend exclusively to the lit arm.We believe the best explanation of these data is that alcohol-treated rats experienced a decrease in attentional capacity andshifted their attention towards the most salient environmentalstimulus, i.e. the lit arm. It is important to note that in thesecond study, alcohol did not cause any increase in impulsivityas measured by choice of the immediate reward. Ifdisinhibition theory were true, alcohol should have increasedthe impulsivity by increasing the choice of the small,immediate reward regardless of the light.

When differential salience between cues is not an issue,studies have found that alcohol does not increase impulsivity.For example, Evenden (1999) measured reflection-impulsivity by training rats to wait for a light indicatingwhich lever is the correct one to press in order to receive afood pellet. The light signal went on three times. The first lightprovided an unreliable indication of the correct lever, thesecond light was more reliable, and the third light was a very

reliable indication of the correct lever. Thus, a slow responseto the light would enable the rat to have the highest accuracyrate and consequently obtain the highest number of foodpellets. Here, the salient cue was the indicator light, which didnot differ over time. Consequently, no effects of alcohol onimpulsivity were found: the rats accuracy rate remainedunaffected by the administration of alcohol (see also Evenden,1998).

Feola et al. (2000) tested impulsivity using a GoStop task,where animals must respond as quickly as possible to a gostimulus, but must withhold their response to a stop stimuluswhich follows the go stimulus. Impulsivity was measured bythe stop time, or how long it takes for the animal to stopresponding. They found that alcohol increased stop time,suggesting that disinhibition occurred. However, the gostimulus was a visual cue (a light) and the stop stimulus wasan auditory cue (a tone). It is difficult to say, then, whether oneof these cues was more salient than the other. It would beinteresting to know whether the same effect would occur if thestimuli were reversed. To include tests of alcohol myopia in astop-go task paradigm, it would be important to manipulatethe salience of the stimuli, such as the brightness of a lightor the loudness of a tone. If alcohol myopia is correct,intoxicated animals behaviour will depend more on which-ever cue is more salient (i.e. alcohol-treated animals will goor stop faster when the stimuli are especially salient), butthe behaviour of control animals will depend less on thesalient cue. Alternatively, counterbalancing the stimulisignalling go and stop would be helpful, as this wouldensure that the results were not due to the specific types ofstimuli used.

To sum up, when impulsivity is measured using the delay-of-reward paradigm or with the gostop task, alcohol oftenappears to increase impulsivity. However, in the Evendenstudies (1998, 1999), low and moderate doses of alcoholappear to have no influence on impulsivity. In the Evendenstudies, the salience of the cue (i.e. the indicator light) was heldconstant. Alcohol myopia would indeed predict that alcoholshould have no effect on impulsivity in such a paradigm. Withthe exception of the study by P. C. Darling, T. L. Pinder,K. G. C. Hellemans, T. A. Paine and M. C. Olmstead (2003)(unpublished data), most studies were not designed in such away that alcohol myopia could be tested. Therefore, it isnot clear whether the increased impulsivity found in manystudies is the result of disinhibition or because of the effects ofalcohol myopia.

OTHER BEHAVIOURS

Alcohol myopia could also be tested in lever pressing for foodreward paradigms. To do this, cues could be made impellingor inhibiting by letting the cues signal reward for leverpressing (e.g. presence of green light signals that reward willbe delivered) or no reward for lever pressing (e.g. presence ofred light signals that reward will not be delivered). Followingthis, both cues could be presented simultaneously (e.g. presentboth green and red lights). The salience (e.g. brightness) ofthese cues could also be manipulated. If alcohol myopia weretrue, when conflicting cues are present, alcohol-treatedanimals should respond more to whichever cue is most salient,

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whether it be impelling or inhibiting. If disinhibition theorywere true, alcohol-treated animals should have difficultywithholding the lever-pressing behaviour and will producemore lever pressing in all conditions.

Finally, discriminative stimulus control is another area inwhich alcohol myopia could be tested. One discriminationprocedure that could be used is the matching to sample taskusing compound stimuli. In this task, animals are presented witha compound sample stimulus (e.g. a red patch above horizontallines). They are then presented with the test stimuli (e.g. red andorange patches, or horizontal and vertical lines) and arereinforced if they respond to the stimulus that matches thesample. This task is particularly well suited to testing alcoholmyopia because the compound stimuli should compete forattention. The salience of the stimuli could also be varied.Alcohol myopia would make the counterintuitive prediction thatalcohol treated rats performance would be more accurate thancontrol rats performance when one stimulus is more salientthan another. For example, if rats were presented with a brightred patch together with relatively dim horizontal lines, alcohol-treated rats should attend primarily to the red patch at theexpense of the lines. When they are then presented test stimuliconsisting of red and orange patches, alcohol-treated rats willrespond to the correct patch because they attended primarily tothe red patch in the sample phase. However, control rats shouldbe able to attend to both stimuli simultaneously when presentedwith compound sample stimuli. Therefore, attention to the linesshould diminish attention to the red patch. Control rats wouldthen be less accurate when presented with the test stimuli. Incontrast to alcohol myopia, disinhibition theory would notpredict that alcohol would improve performance on a matchingto sample task.

CONCLUSION

In most animal studies, the salience of environmental cues isnot manipulated, making it difficult to test alcohol myopia.Therefore, when alcohol produces an increase in disinhibitedbehaviour, it is impossible to know whether this is because ofthe disinhibiting effects of alcohol, or whether it is owingto salient impelling environmental cues. We have providedsuggestions for how animal researchers might design studiesto test alcohol myopia in the behavioural domains ofaggression, social behaviours, and impulsivity, among others.We encourage animal researchers to develop ways to testalcohol myopia within animal research paradigms.

Acknowledgements We would like to thank Cella Olmstead and LeeFabrigar for their comments on the earlier drafts of this manuscript. Thisproject was supported by a New Investigator Award from the CanadianInstitutes of Health Research awarded to T.M.

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