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Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation
Open accessAbstract
The motivation to seek social contact may arise from either positive or negative emotional states, as social interaction can be rewarding and social isolation can be aversive. While ventral tegmental area (VTA) dopamine (DA) neurons may mediate social reward, a cellular substrate for the negative affective state of loneliness has remained elusive. Here, we identify a functional role for DA neurons in the dorsal raphe nucleus (DRN), in which we observe synaptic changes following acute social isolation. DRN DA neurons show increased activity upon social contact following isolation, revealed by in vivo calcium imaging. Optogenetic activation of DRN DA neurons increases social preference but causes place avoidance. Furthermore, these neurons are necessary for promoting rebound sociability following an acute period of isolation. Finally, the degree to which these neurons modulate behavior is predicted by social rank, together supporting a role for DRN dopamine neurons in mediating a loneliness-like state. PAPERCLIP.
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TL;DR
Dorsal raphe nucleus (DRN) dopamine neurons undergo synaptic potentiation after 24 hr social isolation, increase activity upon first social contact, and are sufficient and necessary for isolation-induced sociability. Their activation promotes social preference but is aversive without a social target, suggesting they represent a loneliness-like state modulated by social rank.
Key findings
- 24 hr social isolation increased the AMPAR/NMDAR ratio in DRN DA neurons but not VTA DA neurons; isolated mice also showed a greater AMPAR rectification index and NASPM sensitivity, suggesting a relative increase in GluR2-lacking AMPARs.
- In vivo fiber photometry showed that socially isolated mice had a significantly greater DRN DA calcium signal in response to first contact with a juvenile mouse than group-housed mice, and greater than to a novel object.
- Optogenetic activation of DRN DA neurons increased social preference in the three-chamber task but did not support ICSS and produced real-time place avoidance and conditioned place avoidance; photoinhibition reduced social preference after isolation but not in group-housed mice.
- Social rank predicted behavioral effects: dominance positively correlated with activation-induced social preference and negatively with stimulation-zone preference, while photoinhibition reduced social preference more in dominant mice after isolation. DRN DA neurons also released DA and glutamate in dlBNST and CeA; EPSCs were elicited in 25/30 dlBNST neurons and 17/21 CeA neurons, and 77.7% of eYFP+ DRN neurons in TH::Cre mice and 79.8% in DAT::IRES-Cre mice co-labeled with TH.
Methods
- Ex vivo whole-cell patch-clamp electrophysiology in brain slices (220 μm) from male TH-GFP or Pitx3-GFP mice, measuring AMPAR/NMDAR ratio, rectification index, paired-pulse ratio, and NASPM sensitivity after 24 hr social isolation versus group housing.
- Fiber photometry in TH::Cre mice with AAV5-CAG-FLEX-GCaMP6m targeted to DRN DA neurons; GCaMP6m fluorescence was recorded in the home cage before and after introduction of a juvenile mouse or novel object.
- Optogenetic and behavioral experiments in TH::Cre and DAT::Cre mice with AAV5-DIO-ChR2-eYFP, AAV5-DIO-NpHR3.0-eYFP, or AAV5-DIO-eYFP in DRN; assays included three-chamber sociability, ICSS, real-time place avoidance, conditioned place preference, in vivo fast-scan cyclic voltammetry, and immunohistochemistry.
Limitations
- The paper states that “loneliness” per se is difficult to directly test in mice; it also states that it remains undetermined whether underlying neural differences cause social rank and/or whether social rank itself changes neuronal properties, and that CeA DA transients were inconsistent and did not adequately resolve as DA.
- Evident from the text: ex vivo electrophysiology used male mice, and dominance measures are described for males, so sex differences are not addressed.
- Evident from the text: social-rank effects are correlational, and the “loneliness-like state” interpretation is inferred from mouse behavior rather than directly measured subjective experience.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Finally, the degree to which these neurons modulate behavior is predicted by social rank, together supporting a role for DRN dopamine neurons in mediating a loneliness-like state.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Abstract
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In the paper
“DRN DA neurons show increased activity upon social contact following isolation, revealed by in vivo calcium imaging.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Abstract
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In the paper
“Optogenetic activation of DRN DA neurons increases social preference but causes place avoidance.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Abstract
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In the paper
“Furthermore, these neurons are necessary for promoting rebound sociability following an acute period of isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Abstract
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In the paper
“Here, we identify a functional role for DA neurons in the dorsal raphe nucleus (DRN), in which we observe synaptic changes following acute social isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Abstract
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In the paper
“Optogenetic activation of DRN DA neurons increases social preference but causes place avoidance.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Abstract
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In the paper
“We observed particularly dense terminal expression within the dorsolateral BNST (dlBNST) (Figure 3E) and the lateral part of the CeA (Figure 3F) and, therefore, tested the effects of optical activation of DRN DA neurons on these regions.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“However, in 3/23 dlBNST neurons and 4/24 CeA neurons we observed IPSC responses with a long and variable latency (Figures 4G and 4H), suggesting that terminal stimulation can activate GABAergic neurons to elicit a polysynaptic IPSC.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“In TH::Cre mice we found that 77.7% of eYFP+ neurons co-labeled with TH using immunohistochemistry (Figures S2B, S2D, and S2F).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“Optical stimulation of DRN DA neurons elicited DA release in both the dlBNST (Figure 3G) and the CeA (Figure 3H).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“While eYFP-expressing mice showed the typical trend toward an increase in social preference following isolation, optical inhibition in NpHR-expressing mice resulted in a significant decrease in social preference following isolation (Figures 6D and 6E).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Photoinhibition of DRN DA Neurons Reduces Isolation-Induced Sociability
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In the paper
“There was no significant difference in the proportion of neurons responding with an EPSC in TH::Cre and DAT::Cre mice, so these data were pooled (Figures S4E–S4G).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“The peak-evoked DA release was greater in the dlBNST than the CeA at 30 Hz and 50 Hz (Figures S3D–S3F), suggesting possible differences in the dynamics of DA release and reuptake in these two regions.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“We found that the rectification index (RI) of the AMPAR current was significantly greater in socially isolated mice (Figure 1D), suggesting an increase in GluR2-lacking AMPARs, which exhibit a characteristic inwardly rectifying current at positive potentials (Bellone and Lüscher, 2006, Liu and Zukin, 2007).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Social Isolation Changes Receptor Composition at Synapses onto DRN DA Neurons
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In the paper
“Strikingly, in socially isolated mice, we observed a significant increase in the fluorescence signal in response to first contact with the juvenile mouse, compared with group-housed mice (Figures 2C–2F; Movie S1).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Activity of DRN DA Neurons In Vivo Increases upon Initial Social Contact after Social Isolation
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In the paper
“Consistent with previous reports, eYFP expression in DRN DA somata resulted in terminal expression in several regions including the medial prefrontal cortex (mPFC), bed nucleus of the stria terminalis (BNST), lateral hypothalamus, central amygdala (CeA), entorhinal cortex, and basolateral amygdala (Hasue and Shammah-Lagnado, 2002, Meloni et al., 2006, Swanson, 1982, Yoshida et al., 1989).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
What the sources say
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Furthermore, we did not observe significant effects of optical inhibition on behavioral measures of arousal or anxiety-related behavior (Figures S6B–S6J).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Photoinhibition of DRN DA Neurons Reduces Isolation-Induced Sociability
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In the paper
“Indeed, we found that the paired-pulse ratio (PPR) in the presence of spermine was greater in socially isolated mice (Figures S1F and S1G).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Social Isolation Changes Receptor Composition at Synapses onto DRN DA Neurons
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In the paper
“Furthermore, the EPSCs persisted in the presence of tetradotoxin (TTX) and 4-aminopyridine (4AP), suggesting that they represent monosynaptic glutamate release from DRN DA terminals (Petreanu et al., 2007).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“For comparison, we also examined the DRN of DAT::IRES-Cre mice and found a similar proportion (79.8%) of eYFP+ neurons were colabeled with TH (Figure S2C, S2E, and S2G).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“With Cre-dependent expression of eYFP in the DRN, a subset of TH+ neurons co-expressed VGLUT2 (Figures 4I and 4J), yet there was almost no overlap with VGAT (Figures 4K and 4L).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“However, we found that optical stimulation of DRN DA neurons did not support ICSS in TH::Cre (Figures 5E and 5F) or DAT::Cre mice (Figure S5E).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
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In the paper
“Here, we observed avoidance of the light-paired side of the chamber relative to the unstimulated side in ChR2-, but not eYFP-expressing, TH::Cre and DAT::Cre mice (Figures S5F–S5H).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
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In the paper
“Furthermore, in isolated mice, the activity in response to initial social contact was significantly greater than in response to initial interaction with a novel object (Figures 2C–2F).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Activity of DRN DA Neurons In Vivo Increases upon Initial Social Contact after Social Isolation
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In the paper
“We found that in TH::Cre mice expressing ChR2, but not eYFP, optical stimulation resulted in a significant increase in the proportion of time spent in the social zone (Figures 5A–5D), a result we replicated in DAT::Cre mice (Figures S5C and S5D).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
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In the paper
“Contrastingly, we did not detect a significant difference in decay time constant of the NMDAR current between group-housed and socially-isolated mice (Figure S1H).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Social Isolation Changes Receptor Composition at Synapses onto DRN DA Neurons
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In the paper
“We found that transfer into a new cage as a group had no detectable effect on AMPAR/NMDAR ratio, but social isolation in a new cage also increased the AMPAR/NMDAR ratio (Figure 1C).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Acute Social Isolation Potentiates Synapses onto DA Neurons in the DRN, but Not the VTA
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In the paper
“This revealed that mice expressing NpHR spent a significantly lower proportion of time in the social zone, compared with eYFP-expressing mice (Figure 6C).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Photoinhibition of DRN DA Neurons Reduces Isolation-Induced Sociability
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“With Cre-dependent expression of eYFP in the DRN, a subset of TH+ neurons co-expressed VGLUT2 (Figures 4I and 4J), yet there was almost no overlap with VGAT (Figures 4K and 4L).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“However, we did not detect a difference in AMPAR/NMDAR ratio between group-housed and socially isolated mice in VTA DA neurons (Figure 1A).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Acute Social Isolation Potentiates Synapses onto DA Neurons in the DRN, but Not the VTA
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In the paper
“This reduced the amplitude of the evoked AMPAR current recorded in DRN DA neurons from socially isolated, but not group-housed, mice (Figure 1E).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Social Isolation Changes Receptor Composition at Synapses onto DRN DA Neurons
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In the paper
“Importantly, eYFP expression did not overlap with 5-HT+ (serotonergic) neurons in either TH::Cre or DAT::Cre mice (Figures S2B–S2G).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We found that transfer into a new cage as a group had no detectable effect on AMPAR/NMDAR ratio, but social isolation in a new cage also increased the AMPAR/NMDAR ratio (Figure 1C).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Acute Social Isolation Potentiates Synapses onto DA Neurons in the DRN, but Not the VTA
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We found that in TH::Cre mice expressing ChR2, but not eYFP, optical stimulation resulted in a significant increase in the proportion of time spent in the social zone (Figures 5A–5D), a result we replicated in DAT::Cre mice (Figures S5C and S5D).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
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In the paper
“During the test session, ChR2-expressing mice showed significant avoidance of the stimulation-associated zone, relative to eYFP-expressing mice (Figures 5H–5J).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
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In the paper
“In contrast, optical stimulation of DAergic terminals did not elicit a short-latency GABAA-mediated inhibitory postsynaptic current (IPSC) in the dlBNST or CeA (Figures 4C and 4D).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“Photostimulation of ChR2-expressing DA terminals elicited a short-latency fast AMPAR-mediated excitatory postsynaptic current (EPSC) in 25/30 neurons recorded in the dlBNST and 17/21 neurons in the CeA (Figure 4C-F).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“Strikingly, DRN DA neurons in mice that were socially isolated for 24 hr exhibited a significantly greater AMPAR/NMDAR ratio than group-housed, naive mice (Figure 1C).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Acute Social Isolation Potentiates Synapses onto DA Neurons in the DRN, but Not the VTA
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In the paper
“Intriguingly, we found that relative dominance and change in social preference were positively correlated, such that optical stimulation of DRN DA neurons appeared to be more effective at promoting social preference in dominant mice (Figures 7C, 7D, and S7C).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Prior Social Experience Predicts Functional Role of DRN DA Neurons
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes- Rests on an assumption: The subjective experience of social interaction differs between members of a social group depending on their social rank.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In response to eight pulses of 30 Hz stimulation, delivered every 5 s, DA transients were consistently recorded in the dlBNST.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
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In the paper
“in group-housed mice, we observed limited changes in DRN activity related to initial juvenile contact (Figure 2)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
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In the paper
“Conversely, we observed a significant negative correlation (Figures 7G, 7H, S7E, and S7F), indicating that photoinhibition produced a greater reduction in social preference following isolation in more dominant mice.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Prior Social Experience Predicts Functional Role of DRN DA Neurons
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes- Rests on an assumption: The subjective experience of social interaction differs between members of a social group depending on their social rank.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“we also demonstrate that glutamate can be released with optical stimulation of the DRN DA neurons”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
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In the paper
“following social isolation, photoinhibition caused a reduction in social preference (Figure 6).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
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In the paper
“Our finding that acute social isolation induces synaptic plasticity at synapses onto DRN DA neurons (Figure 1)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
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In the paper
“With photoactivation, we revealed that activating DRN DA neurons promoted social preference in group-housed mice (Figure 5).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“photoinhibition of these neurons did not alter social behavior (Figure 6).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Here, we observed a negative correlation between relative dominance and preference for the stimulation zone, such that more dominant mice displayed greater avoidance of the light-paired side of the chamber (Figures 7E, 7F, and S7D).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Prior Social Experience Predicts Functional Role of DRN DA Neurons
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passes- Rests on an assumption: The subjective experience of social interaction differs between members of a social group depending on their social rank.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“However, in the absence of a social target, mice avoided photoactivation of DRN DA neurons”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesThe paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Therefore, this may be one reason why manipulations of DRN DA activity were not as effective in promoting behavioral adaptations in subordinate animals.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Consistently, in socially isolated mice, we observed a significantly greater increase in DRN DA activity during initial contact with a social target (Figure 2), compared with group-housed mice.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“we observed a relationship in which dominant mice were more sensitive to the behavioral effects of manipulating DRN DA activity (Figure 7).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This suggests that activation of DRN DA neurons plays a causal role in driving social behavior.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This supports the hypothesis that DRN DA activity is required for motivating sociability in response to the negative state of isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Photoinhibition of DRN DA Neurons Reduces Isolation-Induced Sociability
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“Therefore, the negative state of isolation can trigger the motivation to seek and engage in social contact (Baumeister and Leary, 1995, Maner et al., 2007, Williams and Sommer, 1997), perhaps as an evolutionarily conserved mechanism to maintain social connections (Buss, 1990).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“Conversely, the absence of social contact also triggers a strong desire to seek social interaction.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“We found that the rectification index (RI) of the AMPAR current was significantly greater in socially isolated mice (Figure 1D), suggesting an increase in GluR2-lacking AMPARs, which exhibit a characteristic inwardly rectifying current at positive potentials (Bellone and Lüscher, 2006, Liu and Zukin, 2007).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Social Isolation Changes Receptor Composition at Synapses onto DRN DA Neurons
What the sources say
- Not checked yetcites Bellone and Lüscher, 2006
- Not checked yetcites Liu and Zukin, 2007
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Furthermore, the EPSCs persisted in the presence of tetradotoxin (TTX) and 4-aminopyridine (4AP), suggesting that they represent monosynaptic glutamate release from DRN DA terminals (Petreanu et al., 2007).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
What the sources say
- Not checked yetcites Petreanu et al., 2007 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“Social isolation is also aversive to rodents.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This suggests that prior social environment contributes to subsequent isolation-induced synaptic strength.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Prior Social Experience Predicts Functional Role of DRN DA Neurons
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“Amid the sparse existing knowledge of the functional role of DRN DA neurons, optical stimulation of these neurons does not support intra-cranial self-stimulation (ICSS) (McDevitt et al., 2014), in contrast to the VTA (Witten et al., 2011), suggesting that DRN and VTA DA neurons may be functionally distinct.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This suggests that, following social isolation, the presence of a social stimulus is associated with a significant increase in DRN DA activity in vivo.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Activity of DRN DA Neurons In Vivo Increases upon Initial Social Contact after Social Isolation
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Taken together, this suggests that social isolation induces a relative increase in GluR2-lacking AMPARs at glutamatergic synapses onto DRN DA neurons (Figure 1F).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Social Isolation Changes Receptor Composition at Synapses onto DRN DA Neurons
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“Even an acute period of isolation in rodents increases motivation to seek out and engage with conspecifics (Niesink and van Ree, 1982, Panksepp and Beatty, 1980).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
- Not checked yetcites Niesink and van Ree, 1982
- Not checked yetcites Panksepp and Beatty, 1980
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Collectively, these data suggest that inhibition of DRN DA neurons prevents the typical restoration of social contact following a period of isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Photoinhibition of DRN DA Neurons Reduces Isolation-Induced Sociability
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Taken together, this demonstrates that optical stimulation of ChR2-expressing DRN DA neurons is sufficient to trigger rapid DA and glutamate release in two major downstream targets.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“However, in 3/23 dlBNST neurons and 4/24 CeA neurons we observed IPSC responses with a long and variable latency (Figures 4G and 4H), suggesting that terminal stimulation can activate GABAergic neurons to elicit a polysynaptic IPSC.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
Strength:hedged· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This suggests that, in stark contrast to VTA DA neurons, optical activation of DRN DA neurons produces an aversive state.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Therefore, under “sated” group-housed conditions, this type of motivation may not be playing a major role.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Importantly, this also suggests that DRN DA neurons are not merely indicating the removal of sensory stimuli by social isolation but actually representing the subjective experience of a loneliness-like state.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“the observation that socially isolated and subordinate monkeys show similar D2/3 receptor availability (Morgan et al., 2002)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
What the sources say
- Not checked yetcites Morgan et al., 2002 source
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“In humans, perceived social isolation predicts a poor outcome in numerous physical and mental health-related measures, entirely independent of the level of objective isolation (Adam et al., 2006, Cacioppo et al., 2006, Hawkley et al., 2006, Wen et al., 2006).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
What the sources say
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“The relative bias of the DRN DA neurons in their projections to the BNST and CeA (Hasue and Shammah-Lagnado, 2002, Meloni et al., 2006)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
What the sources say
- Not checked yetcites Hasue and Shammah-Lagnado, 2002
- Not checked yetcites Meloni et al., 2006 source
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“it is likely that both of these regions may be important in mediating the observed effects of DRN DA stimulation on behavior.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
Strength:hedged· not yet calibratedCausal:conditional· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Therefore, in their representation of a loneliness-like state, changes in DRN DA activity may only exert a significant effect on the behavior of individuals who are engaged in positively valued social interactions.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Indeed, 24 hr of social isolation results in a blunting of long-term potentiation (LTP) in the BNST (Conrad et al., 2011).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
What the sources say
- Not checked yetcites Conrad et al., 2011 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIn the paper
“In monkeys, the attainment of a dominant social rank increases striatal D2/3 receptor availability (Morgan et al., 2002)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
What the sources say
- Not checked yetcites Morgan et al., 2002 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“suggesting that stimulation is aversive.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Thus, in all cases, dominant mice showed a greater degree of behavioral modulation upon optogenetic manipulation of DRN DA neurons.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Prior Social Experience Predicts Functional Role of DRN DA Neurons
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes- Rests on an assumption: The subjective experience of social interaction differs between members of a social group depending on their social rank.
In the paper
“The characteristics we have observed in DRN DA neurons bear remarkable similarities to the hypotheses generated from human psychology in describing the “need to belong” (Baumeister and Leary, 1995).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
- Not checked yetcites Baumeister and Leary, 1995 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“more lonely individuals show increased attention toward social cues (Gardner et al., 2005, Pickett et al., 2004).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This indicates that the activity within these neurons may only be necessary in situations in which the motivation for social contact is high, such as that experienced after social isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“supports the notion that subordinate animals may be in a loneliness-like state, even while group-housed.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“represents an important distinction from the VTA population and suggests these DA neurons are part of a distinct circuit.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
Strength:hedged· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesNone of the sources cited for this claim have been checked yet.
In the paper
“in a state of loneliness, wherein an individual’s basic need for social connection is unmet (Peplau, 1978), processing of socially relevant information should be prioritized (Baumeister and Leary, 1995).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
- Not checked yetcites Peplau, 1978
- Not checked yetcites Baumeister and Leary, 1995 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our data present an additional element for consideration in the control of social behavior, and support a novel role for a relatively unstudied population of DA neurons in representing the experience of social isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Conclusion
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“in humans, socially excluded individuals display an enhanced memory for social events (Gardner et al., 2000)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
- Not checked yetcites Gardner et al., 2000 source
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passes
- It may state two separate claims as one. 13
- Probably not a substantial claim. 20
- The quote only partly states this claim. 1
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Consistent with our recording results, neither NpHR- nor eYFP-expressing mice showed a significant difference in the proportion of time spent in the social zone with optical inhibition (Figure 6A and 6B).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Photoinhibition of DRN DA Neurons Reduces Isolation-Induced Sociability
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We compared data from mice that either remained group-housed or experienced social isolation and found that the magnitude of the AMPAR/NMDAR ratio was positively correlated with the number of previous cagemates in socially isolated, but not continually group-housed, mice (Figures 7A and 7B).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Prior Social Experience Predicts Functional Role of DRN DA Neurons
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“However, in the CeA, transients were inconsistent and signals did not adequately resolve as DA, perhaps suggesting DA release just below the FSCV detection threshold (Figures 3G and 3H).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
Strength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Furthermore, using additional behavioral assays, we found that optical stimulation of DRN DA neurons had no detectable effect on locomotion, novelty preference, or anxiety-related behavior (Figures S5I–S5X).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Social interactions can be rewarding and thereby recruit components of the brain’s reward circuitry, including the ventral tegmental area (VTA) dopamine (DA) neurons and the nucleus accumbens (NAc) (Dölen et al., 2013, Gunaydin et al., 2014, Robinson et al., 2002).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Taken together, we propose that DRN DA neurons represent a neural substrate for the subjective experience of social isolation and serve to promote a response to alleviate this aversive state.”
- Also stated in: § Abstract · hedged
“supporting a role for DRN dopamine neurons in mediating a loneliness-like state.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
- Rests on an assumption: The behavioral and neural state induced by acute social isolation in mice corresponds to the subjective experience of loneliness in humans.
- Also stated in: § Abstract · hedged
None of the sources cited for this claim have been checked yet.
In the paper
“The TH+ DRN neurons have been confirmed as DAergic as they express aromatic L-amino decarboxylase (AADC), the enzyme that catalyzes conversion of L-3,4-dihydroxyphenylalanine, the product of TH, to DA (Lu et al., 2006) and the DA transporter (DAT) (Dougalis et al., 2012).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
What the sources say
- Not checked yetcites Lu et al., 2006
- Not checked yetcites Dougalis et al., 2012 source
Load-bearingStrength:boosted· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Furthermore, DRN TH+ neurons lack dopamine-β-hydroxylase, which is necessary to convert DA to norepinephrine (Nagatsu et al., 1979), and do not express 5-hydroxytryptamine (5-HT) (Stratford and Wirtshafter, 1990).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
What the sources say
- Not checked yetcites Nagatsu et al., 1979 source
- Not checked yetcites Stratford and Wirtshafter, 1990
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We posit that this recapitulates a loneliness-like state, in which social approach is driven to alleviate the aversive state associated with social isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
- Rests on an assumption: The behavioral and neural state induced by acute social isolation in mice corresponds to the subjective experience of loneliness in humans.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In sum, we find that increasing activity of DRN DA neurons in group-housed mice promotes social preference but also elicits a negative affective state in the absence of a social target.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > Optogenetic Activation of DRN DA Neurons Mimics a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Rodents are innately social creatures and prefer social rather than isolate housing (Loo et al., 2001).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
- Not checked yetcites Loo et al., 2001 source
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“In the DRN, ChR2-expressing neurons reliably followed these photostimulation parameters (Figure 3D).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Results > DRN DA Neurons Release DA and Glutamate
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesThe quote only partly states this claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Social isolation, social exclusion, or feelings of social disconnection can lead to loneliness, which is a strongly aversive emotional state in humans and detrimental to physical and mental well-being (Cacioppo et al., 2006, Cacioppo et al., 2014, Holt-Lunstad et al., 2010, House et al., 1988).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“Social isolation, social exclusion, or feelings of social disconnection can lead to loneliness, which is a strongly aversive emotional state in humans and detrimental to physical and mental well-being (Cacioppo et al., 2006, Cacioppo et al., 2014, Holt-Lunstad et al., 2010, House et al., 1988).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
Strength:boosted· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“However, since optogenetic activation of VTA DA neurons increases social interaction (Gunaydin et al., 2014) and supports positive reinforcement (Tsai et al., 2009, Witten et al., 2011), they are thought to play a causal role in social reward.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Introduction
What the sources say
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Perceived social isolation (or loneliness) reflects the quality of an individual’s social interactions (Hawkley et al., 2008, Peplau, 1978) rather than their quantity or frequency.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
What the sources say
- Not checked yetcites Hawkley et al., 2008 source
- Not checked yetcites Peplau, 1978
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“the motivation for social re-connection was hypothesized to be “sensitive to satiation patterns” (Baumeister and Leary, 1995).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
- Not checked yetcites Baumeister and Leary, 1995 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably not a substantial claim.
In the paper
“while in mice, altering synaptic efficacy in the mPFC is sufficient to promote a change in social rank (Wang et al., 2011).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
What the sources say
- Not checked yetcites Wang et al., 2011 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“distinct neural circuits are thought to motivate food consumption related to the rewarding value of food (Nieh et al., 2015)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
- Not checked yetcites Nieh et al., 2015 source
Strength:hedged· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“DA receptor signaling in these regions modulates synaptic transmission and activity (Kash et al., 2008, de la Mora et al., 2010).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
What the sources say
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“suggests that these neurons either play a role in detecting or in reconciling the disparity between the current and desired social environment.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“It has been hypothesized that the “need to belong” represents a powerful motivational drive, comparable to the basic need for food in a state of hunger (Baumeister and Leary, 1995).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
- Not checked yetcites Baumeister and Leary, 1995 source
Strength:hedged· not yet calibratedCausal:correlational· not yet calibratedTests✗ Asserted: fails✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The authors may not put it forward as true. The paper’s conclusions may not depend on it.
In the paper
“the neuropeptide vasoactive intestinal peptide (VIP) is co-expressed in a subset of DA neurons (Dougalis et al., 2012).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
What the sources say
- Not checked yetcites Dougalis et al., 2012 source
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“the BNST has been associated with mediating long-term “tonic” behavioral responses to sustained, diffuse, and/or unpredictable threats”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
What the sources say
- Not checked yetcites Davis et al., 2010 source
Strength:hedged· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Thus, DA, glutamate, VIP, or the coordinated activity of these three neurotransmitters/neuromodulators may be important in facilitating the output of the DRN DA neurons.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
Strength:hedged· not yet calibratedCausal:conditional· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This suggests that activation of these neurons may be recapitulating a loneliness-like state, which is marked by a negative-affective state in which the drive to seek social contact is increased.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Thus, in a similar manner, social behavior may be driven by distinct neural circuits when motivated by social reward and when motivated by the punishment of social isolation.”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
Strength:hedged· not yet calibratedCausal:conditional· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“and the need to obtain food to alleviate the negative state of hunger (Chen et al., 2015, Sternson et al., 2013).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“The BNST and the CeA have been implicated in diverse behavioral functions (Davis et al., 2010, Janak and Tye, 2015, Kim et al., 2013)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
What the sources say
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“whereas the CeA is thought to be more important in the rapid, acute response to threatening stimuli (Davis et al., 2010)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Characterizing Components of the DRN DA Circuit
What the sources say
- Not checked yetcites Davis et al., 2010 source
Strength:hedged· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“While “loneliness” per se is difficult to directly test in mice, and consequently, the lack of animal literature on this phenomenon is noted (Cacioppo et al., 2014)”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > The Subjective Experience of Social Isolation
What the sources say
- Not checked yetcites Cacioppo et al., 2014 source
Strength:plain· not yet calibratedTests✓ Asserted: passes✗ Checkable: fails✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. It may be too vague to check against evidence. The paper’s conclusions may not depend on it.
In the paper
“it has been postulated that the motivation for social re-connection should elicit “goal-orientated behavior” (Baumeister and Leary, 1995).”
§ Dorsal Raphe Dopamine Neurons Represent the Experience of Social Isolation > Discussion > Satisfying the Profile of a Neural Substrate for a Loneliness-like State
What the sources say
- Not checked yetcites Baumeister and Leary, 1995 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably not a substantial claim.
The paper relies on this without support.
The behavioral and neural state induced by acute social isolation in mice corresponds to the subjective experience of loneliness in humans.
“Taken together, we propose that DRN DA neurons represent a neural substrate for the subjective experience of social isolation and serve to promote a response to alleviate this aversive state.”
“supporting a role for DRN dopamine neurons in mediating a loneliness-like state.”
The paper relies on this without support.
The subjective experience of social interaction differs between members of a social group depending on their social rank.
“As a result, the subjective experience of social interaction is likely to differ between members of a social group dependent on their social rank.”
The paper relies on this without support.
Social rank offers a useful estimate of an individual's subjective social experience.
“social rank offers a useful estimate of an individual’s subjective social experience.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
In the citing paper
“Notably, social rank also predicted the magnitude of behavioral effects elicited upon DRN dopamine manipulations in mice118.”
From Neural mechanisms of social homeostasis (2019) · cites this paper as [118]
In the citing paper
“However, the behavioral change observed in these assays was greater in dominant animals relative to subordinates118.”
From Neural mechanisms of social homeostasis (2019) · cites this paper as [118]
In the citing paper
“This observation suggests that prior social experience may influence the ability of the DRN dopamine neurons to modulate behavior.”
From Neural mechanisms of social homeostasis (2019) · cites this paper as [118]
In the citing paper
“The DRN dopamine system might be one source of negative motivational drive in response to social deficit118.”
From Neural mechanisms of social homeostasis (2019) · cites this paper as [118]
In the citing paper
“These neurons are also Vglut2+ at neonatal timepoints [13], consistent with functional data showing glutamate co-transmission from PAG/DR DA neurons in the adult [47,48].”
From Classification of midbrain dopamine neurons using single-cell gene expression profiling approaches (2020) · cites this paper as [47]
In the citing paper
“inhibition of DR DA neurons promoted social avoidance only following an experience of social isolation [154].”
From Neural circuits regulating prosocial behaviors (2023) · cites this paper as [154]
In the citing paper
“acute social isolation stress was found to induce synaptic adaptations on DA neurons in the DR, but not in the VTA [154].”
From Neural circuits regulating prosocial behaviors (2023) · cites this paper as [154]
In the citing paper
“it was not observed after a brief 24-hr period of social isolation, as expected from previous work (Matthews et al., 2016; Figure 1—figure supplement 1A, B).”
From Oxytocin neurons mediate the effect of social isolation via the VTA circuits (2022) · cites this paper as Matthews et al., 2016
In the citing paper
“While in rodents, 24 hr of social isolation does not change synaptic strength onto DA neurons of the VTA (Matthews et al., 2016)”
From Oxytocin neurons mediate the effect of social isolation via the VTA circuits (2022) · cites this paper as Matthews et al., 2016
In the citing paper
“Brief periods of acute social isolation have been reported to activate midbrain regions in humans (Tomova et al., 2020) and increase DA neurons’ activity within the dorsal Raphe nucleus (DRN) in mice (Matthews et al., 2016).”
From Oxytocin neurons mediate the effect of social isolation via the VTA circuits (2022) · cites this paper as Matthews et al., 2016
In the citing paper
“Remarkably, acute social isolation during adulthood changes the synaptic strength at excitatory inputs onto DA neurons of the DRN in rodents, leaving synaptic transmission onto DA neurons of the VTA unaltered (Matthews et al., 2016).”
From Oxytocin neurons mediate the effect of social isolation via the VTA circuits (2022) · cites this paper as Matthews et al., 2016
Citations
Cites
The need to belong: Desire for interpersonal attachments as a fundamental human motivation.
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Social Relationships and Mortality Risk: A Meta-analytic Review
Holt-Lunstad, J., Smith, T. B., & Layton, J. B. (2010) · PLoS Medicine · doi:10.1371/journal.pmed.1000316 · Cited by 7,272 · cited 2×
“Social isolation, social exclusion, or feelings of social disconnection can lead to loneliness, which is a strongly aversive emotional state in humans and detrimental to physical and mental well-being (Cacioppo et al., 2006, 2014; Holt-Lunstad et al., 2010; House et al., 1988).”
BackgroundSocial Relationships and Health
House, J. S., Landis, K. R., & Umberson, D. (1988) · Science · doi:10.1126/science.3399889 · Cited by 7,073
Ultrasensitive fluorescent proteins for imaging neuronal activity
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Self-esteem as an interpersonal monitor: The sociometer hypothesis
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From circuits to behaviour in the amygdala
Janak, P. H., & Tye, K. M. (2015) · Nature · doi:10.1038/nature14188 · Cited by 2,011 · cited 2×
“The BNST and the CeA have been implicated in diverse behavioral functions (Davis et al., 2010; Janak and Tye, 2015; Kim et al., 2013), and DA receptor signaling in these regions modulates synaptic transmission and activity (Kash et al., 2008; de la Mora et al., 2010).”
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Moy, S. S., Nadler, J. J., Perez, A., Barbaro, R. P., Johns, J. M., Magnuson, T. R., Piven, J., & Crawley, J. N. (2004) · Genes, Brain and Behavior · doi:10.1111/j.1601-1848.2004.00076.x · Cited by 1,522 · cited 1×
“We assessed social preference utilizing the three-chamber sociability task, in which time spent on the ‘‘social’’ side of the chamber (containing a juvenile mouse under a wire cup) is used as a measure of sociability (Moy et al., 2004; Silverman et al., 2010).”
MethodPhasic vs Sustained Fear in Rats and Humans: Role of the Extended Amygdala in Fear vs Anxiety
Davis, M., Walker, D. L., Miles, L., & Grillon, C. (2010) · Neuropsychopharmacology · doi:10.1038/npp.2009.109 · Cited by 1,513 · cited 2×
“The BNST and the CeA have been implicated in diverse behavioral functions (Davis et al., 2010; Janak and Tye, 2015; Kim et al., 2013), and DA receptor signaling in these regions modulates synaptic transmission and activity (Kash et al., 2008; de la Mora et al., 2010).”
Background
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Guzik, T. J., MOHIDDIN, S. A., Dimarco, A. D., Patel, V., Savvatis, K., Marelli‐Berg, F. M., Madhur, M. S., Tomaszewski, M., Maffia, P., D’Acquisto, F., et al. (2020) · Cardiovascular Research · doi:10.1093/cvr/cvaa106 · Cited by 1,484
Integration of optogenetics with complementary methodologies in systems neuroscience
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Social determinants of health and survival in humans and other animals
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Gating of social reward by oxytocin in the ventral tegmental area
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Loneliness in the Modern Age: An Evolutionary Theory of Loneliness (ETL)
Cacioppo, J. T., & Cacioppo, S. (2018) · Advances in Experimental Social Psychology · doi:10.1016/bs.aesp.2018.03.003 · Cited by 481
Metadata
- Authors
- Matthews, G. A., Nieh, E. H., Vander Weele, C. M., Halbert, S. A., Pradhan, R. V., Yosafat, A. S., Glober, G. F., Izadmehr, E. M., Thomas, R. E., Lacy, G. D., Wildes, C. P., Ungless, M. A., & Tye, K. M.
- Affiliations
- Hammersmith Hospital (GB)
- Massachusetts Institute of Technology (US)
- Corresponding author
- Ungless, Mark A.
- Tye, Kay M.
- ORCID
- Venue
- Cell
- Publisher
- Elsevier BV
- Published
- 2016-02
- Volume
- 164
- Issue
- 4
- Pages
- 617-631
- ISSN
- 0092-8674 (ISSN-L)
- PMC
- PMC4752823
- OpenAlex
- W2269038554
- Semantic Scholar
- 1adbaade5ef3bde98d2bb90790ddf3f098fbafb7
- Cited by
- 402
- References
- 73
- Citations by year
- 2026: 30 · 2025: 47 · 2024: 46 · 2023: 48 · 2022: 36 · 2021: 56 · 2020: 30 · 2019: 34 · 2018: 33 · 2017: 25 · 2016: 17
- FWCI
- 20.38
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Cell (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DSpace@MIT (Massachusetts Institute of Technology) (submittedVersion, cc-by-nc)
- Spiral (Imperial College London) (submittedVersion, other-oa)
- Elsevier eBooks
- Funders
- JPB Foundation
- PIIF
- PNDRF
- JFDP
- Whitehall Foundation
- Klingenstein Foundation
- NARSAD Young Investigator Award
- Alfred P Sloan Foundation
- New York Stem Cell Foundation
- McKnight Foundation
- Whitehead Career Development Chair
- NIMH (R01-MH102441-01)
- NIA (RF1-AG047661-01)
- NIH (DP2-DK-102256-01)
- U.K. Medical Research Council (MC-A654-5QB70)
- The Royal Society
- Simons Center
- NSF Graduate Research Program Fellowship
- Integrative Neuronal Systems Fellowship
- NIGMS (T32 GM007484)
- NSF
- Undergraduate Research Opportunities Program at MIT
- MIT Summer Research Program
- Northeastern Cooperative Education Program
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Life Sciences › Neuroscience › Endocrine and Autonomic Systems › Neuroendocrine regulation and behavior
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Keywords
- dorsal raphe nucleus, dopaminergic neurons, social isolation, loneliness, social reward, social preferences, place avoidance, social rank, synaptic plasticity, calcium imaging, optogenetic stimulation, negative emotions
- MeSH terms
- Animals, Dopamine, Loneliness, Male, Reward, Synapses, Ventral Tegmental Area, Patch-Clamp Techniques, Glutamic Acid, Mice, Dopaminergic Neurons, Optogenetics, Dorsal Raphe Nucleus, In Vitro Techniques
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