what are mirror neurons and how have they been linked to observational learning?

Curr Biol. 2013 Dec 2; 23(23): R1057–R1062.

What Nosotros Know Currently about Mirror Neurons

Abstract

Mirror neurons were discovered over twenty years ago in the ventral premotor region F5 of the macaque monkey. Since their discovery much has been written well-nigh these neurons, both in the scientific literature and in the popular press. They have been proposed to be the neuronal substrate underlying a vast array of different functions. Indeed and then much has been written about mirror neurons that last twelvemonth they were referred to, rightly or wrongly, as "The nigh hyped concept in neuroscience". Here we try to cut through some of this hyperbole and review what is currently known (and not known) about mirror neurons.

Chief Text

Introduction

Mirror neurons are a class of neuron that modulate their activeness both when an private executes a specific motor act and when they observe the same or similar act performed by some other individual. They were outset reported in the macaque monkey ventral premotor area F5 [1] and were named mirror neurons in a subsequent publication from the same group [2]. Always since their discovery, there has been great interest in mirror neurons and much speculation most their possible functional part with a particular focus on their proposed role in social cognition. As Heyes [3] wrote "[mirror neurons] intrigue both specialists and non-specialists, historic equally a 'revolution' in understanding social behaviour … and 'the driving force' behind 'the great spring forward' in human evolution…". Indeed then much has been written in both peer-review literature and elsewhere about mirror neurons and their proposed functional part(s) that they have recently been given the moniker "The most hyped concept in neuroscience" [4].

For united states of america, the discovery of mirror neurons was exciting because it has led to a new mode of thinking about how we generate our ain deportment and how we monitor and translate the actions of others. This discovery prompted the notion that, from a functional viewpoint, action execution and observation are closely-related processes, and indeed that our ability to interpret the actions of others requires the involvement of our own motor arrangement.

The aim of this article is not to add to this literature on the putative functional role(south) of mirror neurons, but instead to provide a review of the studies that accept direct recorded mirror neuron action. To date, there accept been over 800 published papers on mirror neurons (from a PubMed search using: "mirror neuron" OR "mirror neurons"). Hither, we restrict our attention to only the primary literature on mirror neurons. Mirror neurons were originally defined equally neurons which "discharged both during monkey's active movements and when the monkey observed meaningful hand movements made by the experimenter" [2]. Thus, the key characteristics of mirror neurons are that their activity is modulated both by action execution and action ascertainment, and that this activity shows a caste of action specificity. This distinguishes mirror neurons from other 'motor' or 'sensory' neurons whose discharge is associated with either execution or ascertainment, simply not both. It also distinguishes mirror neuron responses from other types of response to vision of objects or other non-activity stimuli. Equally the activity of mirror neurons cannot yet be unambiguously detected using neuroimaging techniques, we accept excluded homo and not-human primate imaging studies from this review. Nosotros therefore focus on the 25 papers [ane,2,v–27] that have reported quantitative results of recording mirror neurons or mirror-like neurons in macaque monkeys since 1992 (Table 1).

Table 1

Proportion of neurons recorded in macaque premotor cortex (area F5) and posterior parietal cortex that showed mirror neuron properties.

Reference	Recording expanse	No. neurons	No. mirror	% mirror	¹Activity specificity	Observed effector
Bonini et al.[5]	F5	154	36	23.4%	y	Hand
Caggiano et al.[6]	F5	299	149	49.8%	n	Hand
Caggiano et al.[8]	F5	219	105	48%	n	Hand
Caggiano et al.[seven]	F5	224	123	54.9%	due north	Hand (video)
Caggiano et al.[nine]	F5	785	247	31.5%	n	Mitt (video)
Ferrari et al.[eleven]	F5	485	130	26.viii%	y	Rima oris
Ferrari et al.[12]	F5	209	52	24.nine%	y	Hand
Gallese et al.[2]	F5	532	92	17.three%	y	Mitt
Kohler et al.[16]ⁱⁱ	F5	497	63	12.7%	y	Auditory
Kraskov et al.[17]	F5	64	31	48.four%	y	Mitt (PTNs)
di Pellegrino et al.[1]	F5	184	18	9.8%	y	Manus
Rizzolatti et al.[18]	F5	300	sixty	20%	y	Mitt
Rochat et al.[19]	F5	282	92	32.6%	y	Mitt
Umilta et al.[23]	F5	220	103	46.eight%	y	Hand
Bonini et al.[v]	IPL	120	28	23.3%	y	Hand
Fogassi et al.[13]	IPL	165	41	24.8%	y	Hand
Rozzi et al.[20]	IPL	423	51	12%	y	Manus
Shepherd et al.[21]	LIP	153	30	19.6%	northward	Eye-gaze
Dushanova and Donoghue [x]	M1	303	105	34.half dozen%	y	Reaching
Tkach et al.[22]	M1	829	581	seventy.i%	y	Tracking arm
Vigneswaran et al.[24]	M1	132	77	58.3%	n	Mitt (PTNs)
Tkach et al.[22]	PMd	128	77	60.ane%	y	Tracking arm
Ishida et al.[14]	VIP	541	48	8.ix%	y	Bimodal tactile/visual
Fujii et al.[27]	PM³	148	_	3–14%^iv	n	Mitt
	IPS⁵	148	-	10–42%^iv	n

Mirror neurons were offset described in the rostral division of the ventral premotor cortex (expanse F5) of the macaque brain, and accept subsequently been reported in the inferior parietal lobule, including the lateral and ventral intraparietal areas, and in the dorsal premotor and primary motor cortex. But despite the large array of areas in which mirror neurons accept been reported, the bulk of mirror neuron research has studied the activity of mirror neurons in area F5 (15/25 papers;Figure 1A).

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Number of mirror neurons recorded in areas F5 and in the IPL.

(A) The percentage of mirror neurons every bit a function of publication year for studies reporting mirror neurons in F5 when observing mitt actions. The black line shows the line of best fit. (B) The per centum of mirror neurons in premotor expanse F5 and in the inferior parietal lobule (IPL). The boilerplate per centum of mirror neurons for each region is shown in blackness and the percent of total mirror neurons is shown in gray with the total number of mirror neurons and neurons recorded given above.

Mirror Neurons in Ventral Premotor Region F5

Of the 15 papers reporting mirror neuron activity in area F5, eleven provide details of the number of mirror neurons recorded when observing the experimenter (not a video) reaching and grasping objects with their manus. On average, 33.6% of neurons recorded in F5 have been described equally mirror neurons when the monkey observed hand deportment performed by a human experimenter in forepart of them (ranging from 9.eight–49.eight%; Effigy iA,B). Information technology is of note that the percentage of mirror neurons reported appears to increase as a function of fourth dimension. This most likely reflects a sampling bias during information collection.

The outset three papers [one,2,18] described the bones properties of mirror neurons, and their percentages are depression compared with later studies. The more recent papers, in full general, have investigated modulations of mirror neuron activity with some form of task manipulation. The methodological approach of these afterward papers is to outset select neurons based on their motor properties (for example, selectivity for grasping) and then investigate the responses of this neuronal population to observed actions. This subtle change in the experimental strategy might explicate the apparent increase in the per centum of mirror neurons in F5 equally a function of fourth dimension. Some investigators have avoided the sampling bias based on mirror properties by studying identified pyramidal tract neurons in expanse F5, selected on the basis of their antidromic response and not for their properties during action execution or ascertainment [18]. A large proportion of pyramidal tract neurons in F5 and in M1 appear to prove mirror-like responses (Table 1).

The three early papers [one,2,xviii] provided details about the relative selectivity of mirror neuron belch during action execution and observation. On boilerplate, 48.9% of mirror neurons were classified as broadly congruent. Some mirror neurons discharged for only one action type, such as grasping, during both execution and observation, but showed no specificity for the type of grasp, for example precision grip or whole hand prehension. Others discharged for more than ane type of observed activity, for example grasping and holding. One of the three papers [2] describes a farther category of mirror neurons, strictly congruent mirror neurons; these are defined as mirror neurons that answer selectively to one action type, such equally precision grip, during both activeness execution and observation, and are reported as constituting 31.5% of mirror neurons recorded. Ii of the three papers [ii,18] report a farther category of neuron in F5 that discharged during action observation but non during action execution; on boilerplate these neurons, which would non exist included equally mirror neurons, have been reported as making upward 5.1% of the neurons in F5.

Further neuroanatomical studies of area F5 have revealed three interconnected sub-divisions [28]. The sub-division in which mirror neurons are located is suggested to be on the convexity of the precentral gyrus, side by side to the junior limb of the arcuate sulcus, and referred to every bit surface area F5c. This is distinguished from area F5p (posterior), which is reciprocally connected both with posterior parietal area AIP and primary motor cortex M1, and from area F5a (anterior) in the depth of the sulcus, which has prefrontal connections [29].

Two studies [7,nine] take been reported that have shown that F5 mirror neurons discharged both to the observation of an action performed in forepart of the monkey past the experimenter and to videos of the same action. On boilerplate 26.9% of F5 neurons discharged when the monkey observed a video of a grasping activeness. One of the two studies [7] reported the relative number of mirror neurons that discharged to real and to videoed actions: 46.iv% of neurons in F5 that responded to an executed action too responded when observing a real action, whereas simply 22.3% responded when observing a videoed activeness. Although fewer mirror neurons responded when the monkey was observing the video of an activeness, for those mirror neurons that did discharge, there was no significant departure in the pattern or rate of mirror neuron belch between real and videoed deportment.

2 of the early papers [two,18] on mirror neurons reported that they could not notice any neurons that discharged when monkeys observed an object beingness grasped with a tool. Subsequently, ii studies [12,xix] showed that mirror neurons did respond to such a tool-based activity. In both these latter cases, however, the monkeys had received a loftier exposure to tool use during the grooming catamenia prior to the recordings. One study [12] reported that twenty% of F5 neurons were tool-responding mirror neurons, whereas the other reported the much college percentage of 66.half dozen% [20]. This high percentage most probable reflects a combination of a small sample size (north = 27) and strict inclusion criteria.

Two papers [15,16] have reported that neurons in F5 responded to the sound of an action: then-called auditory mirror neurons. On boilerplate, 17% of F5 neurons have been reported to have auditory properties (12.7% and 21.3%, respectively, in the two papers). 4 papers [vi–8,23] accept reported that mirror neurons not only discharged during activity observation but that their firing is further modulated past different factors: occlusion [23], relative distance of observed action [viii], advantage value [6] and the view point of the observed action [7]. Umilta et al. [23] showed that 19/37 mirror neurons discharged even when the observed activity was occluded or hidden from the observer, demonstrating that direct vision of the action was not necessary to elicit mirror neuron discharge. Caggiano et al. [vii] showed that 149/201 mirror neurons discharged preferentially for 1 or more of three different views of the same action (at 0, xc and 180 degrees). Sixty of these neurons showed a preference for only one view betoken.

Caggiano et al. [8] as well found that F5 mirror neurons have a preference for whether an observed action occurred in peripersonal or extrapersonal infinite: 27/105 mirror neurons discharged preferentially when the observed activeness occurred in the monkeys extra-personal space, whereas 28/105 mirror neurons discharged preferentially when the observed activeness occurred in the monkey's peri-personal space. The remaining 50 mirror neurons showed no preference. Caggiano et al. [6] reported that mirror neuron discharge is modulated by the value of the advantage associated with the action: they showed that 40/87 mirror neurons responded more when a rewarded object was grasped, while xi/87 responded more when observing an action to a not-rewarded action. The remaining mirror neurons showed no preference.

One study [17] recorded from 64 neurons in F5 that were identified as pyramidal tract neurons. Thirty-one of these neurons were classified every bit mirror neurons, with fourteen/31 mirror neurons showing the 'classic' facilitation response during the activeness observation status. Compared with baseline, the action of the remaining 17 mirror neurons was significantly suppressed during action ascertainment. The inclusion of these 'suppression mirror neurons' [viii,17,24,25] clearly changes the overall proportion of neurons responsive during activeness observation.

In a recent study, Maranesi et al. [30] compared multiunit activity responses in areas F5, F4 (premotor regions) and F1 (primary motor cortex, M1). They reported a higher proportion of recording sites showing mirror type responses in area F5 (specially in area F5c), compared with area F4 (caudal role of the ventral premotor cortex) and with F1. In improver, they reported that in penetration sites where they identified mirror responses, they were rarely able to evoke motion using intracortical microsimulation and argued that this might be due to presence of suppression mirror neurons, as showtime identified by Kraskov et al. [17].

One interesting study [27] looked at action in premotor and parietal cortex neurons of the left hemisphere of a Japanese macaque monkey, either while it observed another monkey sitting reverse making reach-to-grasp movements for nutrient rewards, or when it performed similar actions itself. Many neurons in both cortical areas were active during the other monkey's movements, with the proportion varying across different deportment (Table 1). Premotor cortex neurons showed a distinct preference for movements involving the observed monkey'south right arm and hand, and showed a similar preference for the monkey'south own right-sided deportment.

Mirror Neurons in the Junior Parietal Lobule

Iv papers [five,13,20,25] have reported neuronal activity recorded in the inferior parietal lobule that the authors have described as that of mirror neurons (Effigy 1B). None of these papers explicitly specifies the percentage of neurons that were classified as mirror neurons; for 3 of these papers, yet, we were able to judge from the numbers in the papers that the boilerplate percentage of sampled neurons that were mirror neurons was 20% (41/165 Fogassi et al. [13]; 28/120 Bonnini et al. [five]; 51/423 Rozzi et al. [20]).

2 papers [v,13] describe the modulation of mirror neuron activeness in the inferior parietal lobule by the overall goal of the observed action. Here monkeys observed an experimenter reaching for and grasping an object and either placing it in the mouth (eating) or placing it in a container (placing). On average 53% of mirror neurons had a significantly greater firing rate when the monkey observed the 'eating' compared with the 'placing' condition, 17% had a significantly greater firing charge per unit for 'placing' compared with 'eating'. The remaining 30% showed no difference betwixt the two weather condition. Yamazaki et al. [25] reported examples of mirror neuron activeness in macaque expanse inferior parietal lobe; these neurons responded to the aforementioned activeness carried out in rather unlike contexts, suggesting that they are involved in encoding the 'semantic equivalence' of deportment carried out by different agents in different contexts.

Rozzi et al. [20] investigated the properties of mirror neurons in the IPL. They reported that 58% of mirror neurons were responsive to only i type of manus action, for example grasping, and 25% were responsive to two dissimilar paw actions. The remaining 17% were responsive to either observed mouth actions or mouth and hand deportment. Furthermore, they reported that 29% of IPL mirror neurons were strictly coinciding and 54% were broadly coinciding.

Mirror Neurons in the Primary Motor Cortex

The first few papers [2,xviii] that described mirror neurons in area F5 besides reported that the authors found no bear witness of mirror action in M1. Indeed, Gallese et al. [two] argued that, because almost neurons in M1 prove activity during self-motion, the absenteeism of detectable mirror activity in M1 was testify confronting the idea that this activeness might really represent monkey'due south making pocket-size, covert movements while they watched the experimenter. Similarly, a recent multiunit recording study [29] found only a low level of mirror activity within primary motor cortex. Nevertheless, three papers [10,22,24] accept reported mirror neuron-like responses in M1.

Tkach et al. [22] reported that when monkeys either performed a visuomotor tracking task themselves, or watched the same target and cursor being operated by an experimenter, seventy% (581/829) of recorded neurons in M1 showed stable preferred direction tuning during both execution and observation. These authors also reported that 60% (77/128) of neurons in dorsal premotor cortex were modulated in the aforementioned way.

Dushanova and Donoghue [10] recorded from neurons in M1 whilst the monkey either performed a point-to-point arm-reaching task or observed a human experimenter performing the aforementioned action. This report reported that 34.7% (105/303) of the neurons recorded in M1 were directionally tuned during both action execution and activity observation. The mean firing rate during the observation status was on boilerplate 46% of that during the execution condition. In add-on, 38% of neurons retained the same directional tuning during both execution and observation conditions. It should be noted that these studies differ from those previously described that recorded from F5 and IPL.

All the studies on mirror neurons in F5 and IPL accept employed tasks where the macaque monkey observed either a video or the experimenter performing unproblematic achieve and grasp deportment. The two studies [10,22] described above on mirror-like responses in M1 differed in that they used tasks in which the monkey had been extensively trained on the motor execution task. Information technology is unclear whether the relatively high percentage of these mirror-like responses, compared with those in F5 and IPL, reflects differences between the task or real differences in the number of mirror neurons.

The terminal paper [24] on M1 mirror neurons recorded from 132 neurons that were identified as pyramidal tract neurons; 58% of these neurons (77/132) were classified every bit mirror neurons. As in F5, these authors found that these pyramidal tract neurons were either facilitation mirror neurons (58.5%) or suppression mirror neurons (41.five%) during the action ascertainment condition. In dissimilarity to F5, facilitation mirror neurons in M1 fired at significantly lower rates during action observation vs execution, with the former reported as "less than half of that when the monkey performed the grip". It is noteworthy that these authors made simultaneous EMG recordings from upwardly to 11 different arm, hand and digit muscles and confirmed complete absenteeism of activeness during action observation.

Mirror Neurons in Other Regions

Above, we accept described the results of studies reporting mirror neurons in ventral premotor cortex, dorsal premotor cortex, primary motor cortex and inferior parietal lobule. Three further papers [14,21,26] have reported mirror neuron-like responses in two farther areas. The first [14] recorded visuotactile bimodal neurons in the ventral intraparietal area (VIP). These are neurons that showroom tactile receptive fields for a detail body role (for example, face or head) and also exhibit visual receptive fields in the congruent location. This study demonstrated that 48/541 bimodal neurons also exhibited visual receptive fields when observing the congruent expanse being touched on the experimenter. These neurons were not called mirror neurons but 'body-matching bimodal neurons'.

Shepherd et al. [21] reported mirror neuron-similar responses in the lateral intraparietal (LIP) area. These authors reported that thirty/153 neurons in LIP responded not simply when monkeys oriented attending towards the receptive field of those neurons, simply also when they observed other monkeys orienting in the same management.

Yoshida et al. [26] recently recorded from neurons in the medial frontal cortex, some of which selectively responded to cocky or observed deportment within a social context. The neurons were recorded in one of 2 monkeys who, on alternating trials, chose a movement in order to earn a reward. Correct (or incorrect) choices rewarded (or punished: no advantage) both monkeys. 'Partner-type' neurons were selectively responsive to the choices made by the other monkey, signalling the correct or incorrect choice made; interestingly around nineteen% of these 'partner neurons' showed decreased activity during self-movement.

Relating Man Neuroimaging Data to Mirror Neuron Action

Of the over 800 papers returned when searching PubMed for 'mirror neurons' or 'mirror neuron', the vast majority report the results of experiments on human subjects. Of these, the results of human being neuroimaging experiments, specifically fMRI [31], confirm a broad overlap between cortical areas active in humans during activeness observation and areas where mirror neurons take been reported in macaque monkeys (see to a higher place). Thus, changes in the BOLD signal during action observation seem to be consistent with the existence of a mirror neuron system in humans, but they cannot nonetheless furnish conclusive proof. At that place has, however, too been a study of single neuron activity recorded from human neurosurgical patients that has demonstrated mirror neuron activity [32]. Recordings were focused on medial frontal cortex and temporal lobe structures, and show the extensive nature of the mirror neuron system. Unfortunately, neither of the premotor or posterior parietal areas so heavily investigated in monkeys were bachelor for written report in these patients.

Central to being able to interpret human fMRI studies of the mirror neuron organization is understanding the relationship between the BOLD betoken in human and mirror neuron activeness in macaque monkey. To this end, monkey fMRI studies have now demonstrated significant activity during action observation in regions where mirror neurons accept been previously reported [33,34]. These monkey imaging studies have taken advantage of enhancing the neurovascular responses with an fe-based (MION) dissimilarity agent. As with the vast majority of human fMRI studies, however, there is difficulty in relating these results to mirror neurons, in that they only employ an action observation condition and have no action execution status. This makes it difficult to calibrate the action changes in observation to those in execution, and also raises the possibility that sensory responses other than mirror responses contribute to the neurovascular changes (see Introduction).

One possible style of attributing the fMRI response to a single neuronal population, such every bit mirror neurons, is to utilize fMRI adaptation, or repetition suppression. This is a neuroimaging tool that has been adopted to identify neural populations that encode a item stimulus feature [35]. The logic backside fMRI adaptation is that neurons decrease their firing rate with repeated presentations of the stimulus feature that those neurons encode. Past extension information technology has been argued that the BOLD indicate will too decrease with repeated presentations. It has been argued that areas of the cortex that incorporate mirror neurons should testify fMRI adaptation both when an action is executed and afterwards observed, and when an action is observed and subsequently executed. This is considering the stimulus feature encoded in mirror neurons is repeated irrespective of whether the activeness is observed or executed [36].

The results of such studies have produced mixed results. Of the five studies using this technique published to date [36–twoscore], simply iii have demonstrated significant fMRI accommodation consequent with the presence of mirror neurons in the human brain [38–forty]. One possible explanation for the mixed results is that humans exercise have mirror neurons, but that they do not alter their pattern of activation when stimuli that evoke their response are repeated. Indeed a recent written report [9] has shown some evidence that mirror neurons may not modify their firing rate during repetitions of the same activeness; however, in this work the neuronal action represented in the local field potential (LFP) did modulate with repetition. Further work is clearly required to determine why the Assuming signal in humans and the LFP in monkeys practice adapt with repetition, while the evidence to date suggests that mirror neurons may non.

Great intendance must exist taken when comparing the results from man and monkey studies. Specifically, readers must pay careful attention to the difference in the level of inference between the different modalities. The bulk of human being neuroimaging studies written report significant results at the population level where the variance is estimated across subjects. This is in dissimilarity to the studies reporting mirror neurons in macaque monkeys, where the aim is to test whether individual neurons show a consistent modulation of firing charge per unit during periods of action observation and execution. Here the inference is closer to the assay of fMRI at the single subject area level. Therefore, when information technology is reported that xxx% of neurons in any region were significantly modulated during both action observation and execution this does not hateful that the remaining lxx% do not modulate at all. Rather, it means there was not sufficient statistical testify that these neurons displayed mirror activity. Indeed it is quite possible that when tested at the population level, the neurons that are non-significant at the single neuron level could exist significantly modulated when observing an activity.

The point here is that care must be taken when arguing that 'only' X% of neurons in whatever brain region are mirror neurons. The 'only' implies that the remaining neurons are not significantly modulated in any way during action observation. This is not a valid inference as to do and so would exist to accept the zip hypothesis. This may exist peculiarly problematic for cortical regions where responses in individual mirror neurons are relatively weak, such as in M1.

It is ofttimes assumed that mirror neuron activity during action observation is driven, lesser-upwardly, by the visual (or auditory) input. The review of mirror neuron discharge presented hither provides evidence that this is, at all-time, an incomplete description of mirror neuron firing. Nosotros now know that mirror neuron firing rates are modulated by view point [7], value [6] and that they fifty-fifty discharge in the absenteeism of any visual input [23]. This suggests that mirror neurons tin can exist driven or modulated elevation-down past backward connections from other neuronal populations. Indeed, the requirement for such tiptop-down input to regions containing mirror neurons was realized by Jacob and Jeannerod [41], who argued that information technology was incommunicable for a mirror neuron arrangement driven uniquely past the visual input to correctly infer an intention from an observed action if two or more unlike intentions would generate the same activity. The fact that mirror neurons can be driven by astern connections is consistent with recent predictive coding models of mirror neuron office [42–44]. Within this framework, mirror neurons discharge during activity observation not because they are driven by the visual input just considering they are part of a generative model that is predicting the sensory input. This framework provides a theoretical account of mirror neuron activity that resolves the one-to-many mapping problem described by Jacob and Jeannerod [41] and is consistent with top-downwardly modulation of mirror neuron firing rates.

Concluding Remarks

The discovery of mirror neurons has had a profound effect on the field of social cognition. Here we have reviewed what is currently known about mirror neurons in the different cortical areas in which they have been described. At that place is at present evidence that mirror neurons are present throughout the motor system, including ventral and dorsal premotor cortices and primary motor cortex, likewise every bit beingness present in dissimilar regions of the parietal cortex. The functional office(due south) of mirror neurons and whether mirror neurons arise as a result of a functional adaptation and/or of associative learning during development are important questions that still remain to exist solved. In answering these questions nosotros will need to know more than most the connectivity of mirror neurons and their comparative biological science across different species.

Acknowledgements

J.K. and R.N.50. were both funded by the Wellcome Trust, London, UK. We would like to give thanks Alexander Kraskov for helpful comments on an earlier version.

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Source: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3898692/