What Is The Name Of The Neurons That Fire When An Animal Observes And Performs A Goal-oriented Task?

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

What Nosotros Know Currently well-nigh Mirror Neurons

Abstruse

Mirror neurons were discovered over twenty years agone in the ventral premotor region F5 of the macaque monkey. Since their discovery much has been written about 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 final year they were referred to, rightly or wrongly, as "The most 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.

Main Text

Introduction

Mirror neurons are a class of neuron that modulate their activity both when an private executes a specific motor act and when they observe the same or similar act performed past some other individual. They were offset reported in the macaque monkey ventral premotor surface area F5 [1] and were named mirror neurons in a subsequent publication from the same group [two]. Always since their discovery, there has been great interest in mirror neurons and much speculation well-nigh their possible functional part with a particular focus on their proposed function in social cognition. As Heyes [3] wrote "[mirror neurons] intrigue both specialists and non-specialists, celebrated as a 'revolution' in agreement social behaviour … and 'the driving forcefulness' behind 'the bang-up leap forward' in human evolution…". Indeed so much has been written in both peer-review literature and elsewhere virtually mirror neurons and their proposed functional role(s) that they have recently been given the moniker "The nearly hyped concept in neuroscience" [four].

For us, the discovery of mirror neurons was exciting because information technology has led to a new manner of thinking about how nosotros generate our own deportment and how nosotros monitor and translate the deportment of others. This discovery prompted the notion that, from a functional viewpoint, activeness execution and observation are closely-related processes, and indeed that our ability to interpret the actions of others requires the interest of our ain motor system.

The aim of this article is not to add together to this literature on the putative functional function(s) of mirror neurons, but instead to provide a review of the studies that have straight recorded mirror neuron activity. To date, there have been over 800 published papers on mirror neurons (from a PubMed search using: "mirror neuron" OR "mirror neurons"). Here, we restrict our attention to just the main literature on mirror neurons. Mirror neurons were originally defined equally neurons which "discharged both during monkey'south 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 activity execution and activity 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 observation, but not both. Information technology besides distinguishes mirror neuron responses from other types of response to vision of objects or other non-activeness stimuli. As the activity of mirror neurons cannot nonetheless be unambiguously detected using neuroimaging techniques, nosotros take excluded human and non-homo primate imaging studies from this review. We therefore focus on the 25 papers [1,2,5–27] that have reported quantitative results of recording mirror neurons or mirror-like neurons in macaque monkeys since 1992 (Table ane).

Table 1

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

Reference	Recording area	No. neurons	No. mirror	% mirror	1Action specificity	Observed effector
Bonini et al.[v]	F5	154	36	23.4%	y	Hand
Caggiano et al.[6]	F5	299	149	49.8%	n	Paw
Caggiano et al.[viii]	F5	219	105	48%	n	Hand
Caggiano et al.[7]	F5	224	123	54.ix%	n	Hand (video)
Caggiano et al.[9]	F5	785	247	31.5%	n	Hand (video)
Ferrari et al.[11]	F5	485	130	26.8%	y	Mouth
Ferrari et al.[12]	F5	209	52	24.ix%	y	Hand
Gallese et al.[2]	F5	532	92	17.3%	y	Hand
Kohler et al.[16]2	F5	497	63	12.7%	y	Auditory
Kraskov et al.[17]	F5	64	31	48.4%	y	Hand (PTNs)
di Pellegrino et al.[i]	F5	184	xviii	nine.8%	y	Manus
Rizzolatti et al.[18]	F5	300	60	20%	y	Mitt
Rochat et al.[19]	F5	282	92	32.6%	y	Manus
Umilta et al.[23]	F5	220	103	46.8%	y	Mitt
Bonini et al.[5]	IPL	120	28	23.3%	y	Paw
Fogassi et al.[thirteen]	IPL	165	41	24.8%	y	Hand
Rozzi et al.[20]	IPL	423	51	12%	y	Hand
Shepherd et al.[21]	LIP	153	thirty	nineteen.6%	n	Center-gaze
Dushanova and Donoghue [10]	M1	303	105	34.vi%	y	Reaching
Tkach et al.[22]	M1	829	581	lxx.1%	y	Tracking arm
Vigneswaran et al.[24]	M1	132	77	58.3%	due north	Manus (PTNs)
Tkach et al.[22]	PMd	128	77	threescore.1%	y	Tracking arm
Ishida et al.[fourteen]	VIP	541	48	8.nine%	y	Bimodal tactile/visual
Fujii et al.[27]	PM3	148	_	three–xiv%four	n	Mitt
	IPS5	148	-	10–42%four	due north

Mirror neurons were first described in the rostral division of the ventral premotor cortex (area F5) of the macaque brain, and have 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 assortment of areas in which mirror neurons have been reported, the majority of mirror neuron research has studied the activeness of mirror neurons in area F5 (15/25 papers;Effigy 1A).

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

(A) The percentage of mirror neurons as a function of publication twelvemonth for studies reporting mirror neurons in F5 when observing hand actions. The black line shows the line of best fit. (B) The percentage of mirror neurons in premotor expanse F5 and in the inferior parietal lobule (IPL). The average percentage of mirror neurons for each region is shown in black and the percent of full mirror neurons is shown in greyness with the total number of mirror neurons and neurons recorded given above.

Mirror Neurons in Ventral Premotor Region F5

Of the xv papers reporting mirror neuron action in area F5, xi provide details of the number of mirror neurons recorded when observing the experimenter (not a video) reaching and grasping objects with their hand. On average, 33.6% of neurons recorded in F5 take been described every bit mirror neurons when the monkey observed hand actions performed by a man experimenter in front end of them (ranging from 9.8–49.8%; Effigy 1A,B). It is of note that the per centum of mirror neurons reported appears to increase every bit a function of time. This most likely reflects a sampling bias during data collection.

The beginning three papers [1,2,xviii] described the basic properties of mirror neurons, and their percentages are depression compared with later studies. The more contempo papers, in general, have investigated modulations of mirror neuron activity with some form of chore manipulation. The methodological arroyo of these subsequently papers is to first 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 alter in the experimental strategy might explain the apparent increase in the percentage of mirror neurons in F5 equally a role of time. Some investigators have avoided the sampling bias based on mirror properties by studying identified pyramidal tract neurons in area F5, selected on the basis of their antidromic response and not for their properties during action execution or observation [18]. A large proportion of pyramidal tract neurons in F5 and in M1 appear to show mirror-similar responses (Table ane).

The three early on papers [1,2,18] provided details about the relative selectivity of mirror neuron discharge during action execution and observation. On average, 48.ix% of mirror neurons were classified every bit broadly congruent. Some mirror neurons discharged for only one activeness type, such every bit grasping, during both execution and ascertainment, merely showed no specificity for the type of grasp, for example precision grip or whole mitt prehension. Others discharged for more one type of observed action, for example grasping and holding. One of the 3 papers [2] describes a further category of mirror neurons, strictly congruent mirror neurons; these are defined as mirror neurons that respond selectively to ane activity type, such as precision grip, during both action execution and observation, and are reported as constituting 31.5% of mirror neurons recorded. Two of the three papers [ii,18] report a further category of neuron in F5 that discharged during action observation but non during action execution; on boilerplate these neurons, which would not be included as mirror neurons, have been reported equally making up 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, adjacent to the inferior limb of the arcuate sulcus, and referred to every bit area F5c. This is distinguished from area F5p (posterior), which is reciprocally continued both with posterior parietal surface area AIP and main motor cortex M1, and from area F5a (anterior) in the depth of the sulcus, which has prefrontal connections [29].

Two studies [seven,ix] have been reported that have shown that F5 mirror neurons discharged both to the observation of an action performed in front of the monkey by the experimenter and to videos of the aforementioned action. On average 26.9% of F5 neurons discharged when the monkey observed a video of a grasping activity. One of the ii studies [7] reported the relative number of mirror neurons that discharged to real and to videoed deportment: 46.four% of neurons in F5 that responded to an executed activeness also responded when observing a existent action, whereas simply 22.3% responded when observing a videoed action. Although fewer mirror neurons responded when the monkey was observing the video of an action, for those mirror neurons that did discharge, there was no significant deviation in the pattern or rate of mirror neuron belch between real and videoed deportment.

Two of the early on papers [two,18] on mirror neurons reported that they could non find any neurons that discharged when monkeys observed an object existence grasped with a tool. Subsequently, two studies [12,19] showed that mirror neurons did respond to such a tool-based activity. In both these latter cases, however, the monkeys had received a high exposure to tool use during the training period prior to the recordings. One written report [12] reported that 20% of F5 neurons were tool-responding mirror neurons, whereas the other reported the much higher per centum of 66.6% [xx]. This high percentage most probable reflects a combination of a small sample size (n = 27) and strict inclusion criteria.

Two papers [15,xvi] have reported that neurons in F5 responded to the audio of an activity: so-chosen 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 [six–8,23] accept reported that mirror neurons non only discharged during action observation but that their firing is further modulated past different factors: apoplexy [23], relative altitude of observed action [eight], reward value [6] and the view point of the observed activity [vii]. Umilta et al. [23] showed that 19/37 mirror neurons discharged even when the observed action was occluded or hidden from the observer, demonstrating that direct vision of the action was not necessary to arm-twist mirror neuron discharge. Caggiano et al. [seven] showed that 149/201 mirror neurons discharged preferentially for one or more of 3 dissimilar views of the same action (at 0, 90 and 180 degrees). Sixty of these neurons showed a preference for only one view signal.

Caggiano et al. [8] as well plant that F5 mirror neurons have a preference for whether an observed activeness occurred in peripersonal or extrapersonal space: 27/105 mirror neurons discharged preferentially when the observed action occurred in the monkeys extra-personal infinite, whereas 28/105 mirror neurons discharged preferentially when the observed action 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 reward associated with the action: they showed that 40/87 mirror neurons responded more when a rewarded object was grasped, while 11/87 responded more when observing an activity to a non-rewarded activeness. The remaining mirror neurons showed no preference.

1 written report [17] recorded from 64 neurons in F5 that were identified every bit pyramidal tract neurons. Thirty-one of these neurons were classified as mirror neurons, with xiv/31 mirror neurons showing the 'classic' facilitation response during the activity observation status. Compared with baseline, the action of the remaining 17 mirror neurons was significantly suppressed during action observation. The inclusion of these 'suppression mirror neurons' [8,17,24,25] conspicuously changes the overall proportion of neurons responsive during action ascertainment.

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 part of the ventral premotor cortex) and with F1. In addition, they reported that in penetration sites where they identified mirror responses, they were rarely able to evoke movement using intracortical microsimulation and argued that this might be due to presence of suppression mirror neurons, as start identified by Kraskov et al. [17].

One interesting written report [27] looked at activity in premotor and parietal cortex neurons of the left hemisphere of a Japanese macaque monkey, either while it observed another monkey sitting opposite making reach-to-grasp movements for nutrient rewards, or when it performed similar actions itself. Many neurons in both cortical areas were agile during the other monkey's movements, with the proportion varying beyond different actions (Tabular array ane). Premotor cortex neurons showed a distinct preference for movements involving the observed monkey's correct arm and hand, and showed a similar preference for the monkey's own right-sided deportment.

Mirror Neurons in the Inferior Parietal Lobule

Four papers [5,13,20,25] have reported neuronal action recorded in the inferior parietal lobule that the authors have described as that of mirror neurons (Figure 1B). None of these papers explicitly specifies the percentage of neurons that were classified as mirror neurons; for three of these papers, even so, we were able to estimate from the numbers in the papers that the average percent of sampled neurons that were mirror neurons was 20% (41/165 Fogassi et al. [thirteen]; 28/120 Bonnini et al. [5]; 51/423 Rozzi et al. [20]).

Two papers [v,13] describe the modulation of mirror neuron activity 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 oral cavity (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 divergence between the 2 conditions. Yamazaki et al. [25] reported examples of mirror neuron activity in macaque area junior parietal lobe; these neurons responded to the aforementioned action carried out in rather different contexts, suggesting that they are involved in encoding the 'semantic equivalence' of deportment carried out by dissimilar 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 one type of paw activity, for instance grasping, and 25% were responsive to two different hand deportment. 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 congruent and 54% were broadly congruent.

Mirror Neurons in the Primary Motor Cortex

The showtime few papers [2,eighteen] that described mirror neurons in area F5 as well reported that the authors found no evidence of mirror activeness in M1. Indeed, Gallese et al. [2] argued that, because most neurons in M1 bear witness activity during self-movement, the absence of detectable mirror activity in M1 was evidence against the idea that this activity might actually represent monkey'south making small, covert movements while they watched the experimenter. Similarly, a contempo multiunit recording study [29] institute only a depression level of mirror action within primary motor cortex. All the same, three papers [10,22,24] have reported mirror neuron-similar 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, 70% (581/829) of recorded neurons in M1 showed stable preferred management tuning during both execution and ascertainment. These authors also reported that 60% (77/128) of neurons in dorsal premotor cortex were modulated in the same style.

Dushanova and Donoghue [ten] recorded from neurons in M1 whilst the monkey either performed a bespeak-to-point arm-reaching job or observed a human being experimenter performing the aforementioned action. This study reported that 34.vii% (105/303) of the neurons recorded in M1 were directionally tuned during both activeness execution and action observation. The hateful firing rate during the observation status was on average 46% of that during the execution condition. In addition, 38% of neurons retained the aforementioned directional tuning during both execution and observation conditions. Information technology should exist 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 have employed tasks where the macaque monkey observed either a video or the experimenter performing unproblematic reach and grasp actions. The two studies [10,22] described in a higher place on mirror-like responses in M1 differed in that they used tasks in which the monkey had been extensively trained on the motor execution chore. Information technology is unclear whether the relatively high percent of these mirror-like responses, compared with those in F5 and IPL, reflects differences between the job or real differences in the number of mirror neurons.

The final 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 as mirror neurons. Equally in F5, these authors establish that these pyramidal tract neurons were either facilitation mirror neurons (58.5%) or suppression mirror neurons (41.5%) during the activeness observation condition. In contrast to F5, facilitation mirror neurons in M1 fired at significantly lower rates during action ascertainment vs execution, with the former reported every bit "less than one-half of that when the monkey performed the grip". It is noteworthy that these authors fabricated simultaneous EMG recordings from upwardly to 11 different arm, manus and digit muscles and confirmed complete absence of activity during activeness observation.

Mirror Neurons in Other Regions

In a higher place, we have 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 further areas. The first [14] recorded visuotactile bimodal neurons in the ventral intraparietal surface area (VIP). These are neurons that exhibit tactile receptive fields for a particular body part (for instance, confront or head) and also showroom visual receptive fields in the coinciding location. This written report demonstrated that 48/541 bimodal neurons also exhibited visual receptive fields when observing the congruent area beingness touched on the experimenter. These neurons were not called mirror neurons but 'body-matching bimodal neurons'.

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

Yoshida et al. [26] recently recorded from neurons in the medial frontal cortex, some of which selectively responded to self or observed actions within a social context. The neurons were recorded in one of 2 monkeys who, on alternate trials, chose a movement in order to earn a reward. Right (or incorrect) choices rewarded (or punished: no reward) both monkeys. 'Partner-type' neurons were selectively responsive to the choices made by the other monkey, signalling the correct or wrong choice made; interestingly around 19% of these 'partner neurons' showed decreased activity during self-movement.

Relating Human Neuroimaging Data to Mirror Neuron Activeness

Of the over 800 papers returned when searching PubMed for 'mirror neurons' or 'mirror neuron', the vast majority written report the results of experiments on human subjects. Of these, the results of human neuroimaging experiments, specifically fMRI [31], confirm a broad overlap between cortical areas agile in humans during activeness ascertainment and areas where mirror neurons have been reported in macaque monkeys (run into above). Thus, changes in the Assuming signal during action observation seem to exist consequent with the existence of a mirror neuron arrangement in humans, but they cannot yet furnish conclusive proof. There has, nevertheless, as well been a report of single neuron activeness recorded from human neurosurgical patients that has demonstrated mirror neuron activity [32]. Recordings were focused on medial frontal cortex and temporal lobe structures, and prove the extensive nature of the mirror neuron system. Unfortunately, neither of the premotor or posterior parietal areas so heavily investigated in monkeys were available for study in these patients.

Central to beingness able to translate human fMRI studies of the mirror neuron system is understanding the human relationship betwixt the Assuming signal in human and mirror neuron activity in macaque monkey. To this finish, monkey fMRI studies have now demonstrated pregnant activity during action observation in regions where mirror neurons have been previously reported [33,34]. These monkey imaging studies have taken reward of enhancing the neurovascular responses with an iron-based (MION) contrast agent. Equally with the vast majority of human fMRI studies, however, at that place is difficulty in relating these results to mirror neurons, in that they only employ an activity observation condition and accept no action execution condition. This makes it difficult to calibrate the activity 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 as mirror neurons, is to apply fMRI accommodation, or repetition suppression. This is a neuroimaging tool that has been adopted to identify neural populations that encode a particular stimulus characteristic [35]. The logic behind fMRI adaptation is that neurons decrease their firing rate with repeated presentations of the stimulus feature that those neurons encode. Past extension it has been argued that the Assuming betoken will also subtract with repeated presentations. Information technology has been argued that areas of the cortex that contain mirror neurons should testify fMRI accommodation both when an action is executed and later on 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 action is observed or executed [36].

The results of such studies have produced mixed results. Of the five studies using this technique published to appointment [36–xl], only three have demonstrated meaning fMRI accommodation consistent with the presence of mirror neurons in the human encephalon [38–twoscore]. 1 possible explanation for the mixed results is that humans do have mirror neurons, but that they exercise not change their design of activation when stimuli that evoke their response are repeated. Indeed a recent study [ix] has shown some evidence that mirror neurons may not change their firing rate during repetitions of the same action; all the same, in this work the neuronal action represented in the local field potential (LFP) did modulate with repetition. Further piece of work is conspicuously required to determine why the BOLD signal in humans and the LFP in monkeys do adapt with repetition, while the prove to date suggests that mirror neurons may not.

Great intendance must be taken when comparing the results from human being 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 neuroimaging studies report pregnant results at the population level where the variance is estimated across subjects. This is in contrast to the studies reporting mirror neurons in macaque monkeys, where the aim is to examination whether private neurons evidence a consistent modulation of firing rate during periods of action observation and execution. Hither the inference is closer to the analysis of fMRI at the single bailiwick level. Therefore, when it is reported that 30% of neurons in any region were significantly modulated during both action observation and execution this does not mean that the remaining 70% exercise not modulate at all. Rather, it means at that place 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 action.

The signal hither is that intendance must be taken when arguing that 'only' X% of neurons in any brain region are mirror neurons. The 'simply' implies that the remaining neurons are not significantly modulated in whatever manner during action observation. This is not a valid inference every bit to exercise so would be to accept the null hypothesis. This may be particularly problematic for cortical regions where responses in individual mirror neurons are relatively weak, such as in M1.

It is often causeless that mirror neuron activity during action observation is driven, bottom-upwardly, by the visual (or auditory) input. The review of mirror neuron discharge presented hither provides evidence that this is, at best, an incomplete description of mirror neuron firing. Nosotros at present 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 can be driven or modulated height-down by backward connections from other neuronal populations. Indeed, the requirement for such top-downwards input to regions containing mirror neurons was realized by Jacob and Jeannerod [41], who argued that it was incommunicable for a mirror neuron system driven uniquely past the visual input to correctly infer an intention from an observed action if two or more dissimilar intentions would generate the same action. The fact that mirror neurons can be driven by astern connections is consistent with contempo predictive coding models of mirror neuron function [42–44]. Within this framework, mirror neurons discharge during activity ascertainment not considering they are driven by the visual input just because they are part of a generative model that is predicting the sensory input. This framework provides a theoretical business relationship of mirror neuron activity that resolves the one-to-many mapping problem described past Jacob and Jeannerod [41] and is consistent with superlative-downwardly modulation of mirror neuron firing rates.

Concluding Remarks

The discovery of mirror neurons has had a profound consequence on the field of social cognition. Here we have reviewed what is currently known most mirror neurons in the different cortical areas in which they have been described. There is now evidence that mirror neurons are present throughout the motor system, including ventral and dorsal premotor cortices and primary motor cortex, as well as being present in dissimilar regions of the parietal cortex. The functional role(s) of mirror neurons and whether mirror neurons arise as a issue of a functional accommodation and/or of associative learning during development are important questions that even so remain to be solved. In answering these questions nosotros will demand to know more about the connectivity of mirror neurons and their comparative biological science beyond dissimilar species.

Acknowledgements

J.K. and R.N.L. were both funded by the Wellcome Trust, London, Uk. We would like to thank Alexander Kraskov for helpful comments on an before version.

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

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