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ABCDEFGH
1
NeuronTypeLocationDescription
Neurotransmitter/Neuropeptide
Innexin ExpressionReceptor expressionFunction
2
ADAL
Interneuron
Lateral in head, at the level of the second bulb of the pharynx
Integrative ring interneurons- Glutamate
- Possibly FLP-8, a FMRFamide-like peptide
(Loer, 2010; Li and Kim, 2008; Lee et al.,1999)

- INX-4
- INX-19
- UNC-9
(Altun et al, 2009; Chuang et al, 2007)
UnknownUnknown. However, topological cluster analyses suggest ADA may be involved in chemosensation (Sohn et al., 2011).
3
ADAR
Interneuron
Lateral in head, at the level of the second bulb of the pharynx
Integrative ring interneurons- Glutamate
- Possibly FLP-8, a FMRFamide-like peptide
(Loer, 2010; Li and Kim, 2008; Lee et al.,1999)
- INX-4
- INX-19
- UNC-9
(Altun et al, 2009; Chuang et al, 2007
UnknownUnknown. However, topological cluster analyses suggest ADA may be involved in chemosensation (Sohn et al., 2011).
4
ADEL
Sensory neuron
Lateral in head, behind the second bulb of the pharynxAnterior deirids. The sensillar branch from the dorsal process follows a dorsal and frequently posterior trajectory ending laterally within the lateral alae in the anterior third of the body.- Dopamine
(Loer, 2010; Sulston et al., 1975)
- INX-4
- INX-7
- UNC-9
(Altun et al., 2009)
- DOP-2; dopamine receptor
- EXP-1; excitatory, cation-selective GABA receptor
- TYRA-3; tyramine receptor
- Possibly ACR-16; nicotinic acetylcholine receptor
(Altun, 2011; Wragg et al., 2007; Feng et al., 2006; Beg and Jorgensen, 2003; Suo et al., 2003)
- ADEL/R are dopaminergic nose touch mechanoceptors. They modulate locomotion behavior in response to the presence of food by textural mechanosensation, wherein upon encountering a lawn of bacteria (or Sephadex G-200 beads) worms slow their forward movement. The three classes of ciliated dopaminergic neurons (CEPs, ADEs, and PDEs) function redundantly in this behavior (i.e. sensing the mechanosensory stimuli during crawling through bacteria (or the beads) and communicating with the motor circuit to bring about the locomotory change) (Sawin et al., 2000). Another form of motor plasticity related to food presence is area restricted search (ARS). CEP and PDE neurons work synergistically to mediate this behavior while ADE modulates it (Hills et al., 2004).
- Anterior harsh touch sensation: Laser ablation of FLP, AQR, ADE results in a significant reduction of anterior harsh touch sensation, while laser ablation of BDU, SDQR, FLP, ADE and AQR eliminates it (Li et al., 2011).
- Possibly pheromone sensation: The complete pheromone insensitivity of the tax-2(p694) mutant suggests that this defect is due to a site of action in one or more of the AFD, ASE , ADE, or BAG neurons, since the tax-2(p694) mutation eliminates expression of the TAX-2 subunit from these, but has normal expression and function in the other seven neurons that express this channel (Aillion and Thomas, 2000).
5
ADER
Sensory neuron
Lateral in head, behind the second bulb of the pharynxAnterior deirids. The sensillar branch from the dorsal process follows a dorsal and frequently posterior trajectory ending laterally within the lateral alae in the anterior third of the body.- Dopamine
(Loer, 2010; Sulston et al., 1975)
- INX-4
- INX-7
- UNC-9
(Altun et al., 2009)
- DOP-2; dopamine receptor
- EXP-1; excitatory, cation-selective GABA receptor
- TYRA-3; tyramine receptor
- Possibly ACR-16; nicotinic acetylcholine receptor
(Altun, 2011; Wragg et al., 2007; Feng et al., 2006; Beg and Jorgensen, 2003; Suo et al., 2003)
- ADEL/R are dopaminergic nose touch mechanoceptors. They modulate locomotion behavior in response to the presence of food by textural mechanosensation, wherein upon encountering a lawn of bacteria (or Sephadex G-200 beads) worms slow their forward movement. The three classes of ciliated dopaminergic neurons (CEPs, ADEs, and PDEs) function redundantly in this behavior (i.e. sensing the mechanosensory stimuli during crawling through bacteria (or the beads) and communicating with the motor circuit to bring about the locomotory change) (Sawin et al., 2000). Another form of motor plasticity related to food presence is area restricted search (ARS). CEP and PDE neurons work synergistically to mediate this behavior while ADE modulates it (Hills et al., 2004).
- Anterior harsh touch sensation: Laser ablation of FLP, AQR, ADE results in a significant reduction of anterior harsh touch sensation, while laser ablation of BDU, SDQR, FLP, ADE and AQR eliminates it (Li et al., 2011).
- Possibly pheromone sensation: The complete pheromone insensitivity of the tax-2(p694) mutant suggests that this defect is due to a site of action in one or more of the AFD, ASE , ADE, or BAG neurons, since the tax-2(p694) mutation eliminates expression of the TAX-2 subunit from these, but has normal expression and function in the other seven neurons that express this channel (Aillion and Thomas, 2000).
6
ADFL
Sensory neuron
Lateral ganglia in headAmphid neurons, with dual (AdF) ciliated sensory endings which are exposed to the external environment. Enter ring via commissure from ventral ganglion. Take up FITC.- Serotonin
- FLP-6; FMRFamide-like peptide
- INS-1; insulin-like peptide
- NLP-3; neuropeptide-like peptide
(Loer, 2010; Li and Kim, 2008; Kodama et al., 2006; Nathoo et al., 2001; Sze et al., 2000; Duerr et al.,1999)
- INX-4
- INX-19
(Altun et al., 2009; Chuang et al., 2007)
- MGL-3; metabotropic glutamate receptor family protein
- NPR-5; receptor for flp-18 encoded peptides
- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- OCR-2; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- SRB-6 (faint); G protein-coupled seven transmembrane receptor
- SRD-1 (in male); G protein-coupled seven transmembrane receptor
(Altun, 2011; Cohen et al., 2009; Greer et al., 2008; Tobin et al., 2002; Colbert et al., 1997; Troemel et al.,1995)
ADFL/R are the only serotonergic sensory neurons in the worm and may couple environmental food signals with serotonin neurotransmission (Jafari et al., 2011).
- They contribute to a residual chemotactic response to cAMP, biotin, Cl-, and Na+ after ASE is killed (Bargmann and Horvitz, 1991). C. elegans shows chemoattraction to low NaCl. This is mediated by four pairs (ADF, ASE, ASG and ASI) of amphid sensory neurons, of which the ASE cells are most important. It also shows avoidance of NaCl concentrations above 200 mM which is thought to be due to a general avoidance of high osmotic strength (mainly mediated by the ASH sensory neurons). However, this response to salt is plastic, involving a balance between attraction and avoidance, i.e., balances between ASE, ASI, ASH, ADF and perhaps ADL neuron (Hukema et al, 2006).
- Control entry into dauer stage; ADF, ASI and ASG inhibit entry into dauer stage while ASJ and ASK promote dauer entry (Kim et al., 2009; Ouellett et al., 2008; Schackwitz et al., 1996; Bargmann and Horvitz, 1991).
- Function as part of the oxygen-sensing network (which also includes URX, AQR, PQR, SDQ, ALN, PLN, ADL, and ASH) by promoting hyperoxia avoidance. ADF pair stimulates aerotaxis in the absence of food by producing serotonin (Chang et al., 2006).
- Modulate NMJ neurotransmission; serotonin functions as a neuromodulator by inhibiting and enhancing synaptic transmission of other neurotransmitters; e. g. studies have revealed both stimulatory and inhibitory serotonin inputs to the NMJs. Aldicarb, an inhibitor of acetylcholinesterase, causes paralysis in C. elegans due to accumulation of acetylcholine (ACh) at the locomotory NMJs, but exogenous serotonin inhibits paralysis induced by aldicarb. By contrast, exogenous serotonin does not reduce the paralysis induced by levamisole, a specific agonist of the nicotinic ACh receptor UNC-29 in the bodywall muscles. This suggests that serotonin signaling inhibits ACh release by the motor neurons. Studies showed that the inhibitory and stimulatory serotonin signals to NMJ neurotransmission arise from distinct serotonergic neurons; endogenous serotonin released from the ADF neurons stimulates ACh synaptic transmission at the NMJs, whereas serotonin from the NSM/RIH/AIM neurons inhibits the stimulatory serotonin inputs (Govorunova et al., 2010).
7
ADFR
Sensory neuron
Lateral ganglia in headAmphid neurons, with dual (AdF) ciliated sensory endings which are exposed to the external environment. Enter ring via commissure from ventral ganglion. Take up FITC.- Serotonin
- FLP-6; FMRFamide-like peptide
- INS-1; insulin-like peptide
- NLP-3; neuropeptide-like peptide
(Loer, 2010; Li and Kim, 2008; Kodama et al., 2006; Nathoo et al., 2001; Sze et al., 2000; Duerr et al.,1999)
- INX-4
- INX-19
(Altun et al., 2009; Chuang et al., 2007)
- MGL-3; metabotropic glutamate receptor family protein
- NPR-5; receptor for flp-18 encoded peptides
- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- OCR-2; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- SRB-6 (faint); G protein-coupled seven transmembrane receptor
- SRD-1 (in male); G protein-coupled seven transmembrane receptor
(Altun, 2011; Cohen et al., 2009; Greer et al., 2008; Tobin et al., 2002; Colbert et al., 1997; Troemel et al.,1995)
ADFL/R are the only serotonergic sensory neurons in the worm and may couple environmental food signals with serotonin neurotransmission (Jafari et al., 2011).
- They contribute to a residual chemotactic response to cAMP, biotin, Cl-, and Na+ after ASE is killed (Bargmann and Horvitz, 1991). C. elegans shows chemoattraction to low NaCl. This is mediated by four pairs (ADF, ASE, ASG and ASI) of amphid sensory neurons, of which the ASE cells are most important. It also shows avoidance of NaCl concentrations above 200 mM which is thought to be due to a general avoidance of high osmotic strength (mainly mediated by the ASH sensory neurons). However, this response to salt is plastic, involving a balance between attraction and avoidance, i.e., balances between ASE, ASI, ASH, ADF and perhaps ADL neuron (Hukema et al, 2006).
- Control entry into dauer stage; ADF, ASI and ASG inhibit entry into dauer stage while ASJ and ASK promote dauer entry (Kim et al., 2009; Ouellett et al., 2008; Schackwitz et al., 1996; Bargmann and Horvitz, 1991).
- Function as part of the oxygen-sensing network (which also includes URX, AQR, PQR, SDQ, ALN, PLN, ADL, and ASH) by promoting hyperoxia avoidance. ADF pair stimulates aerotaxis in the absence of food by producing serotonin (Chang et al., 2006).
- Modulate NMJ neurotransmission; serotonin functions as a neuromodulator by inhibiting and enhancing synaptic transmission of other neurotransmitters; e. g. studies have revealed both stimulatory and inhibitory serotonin inputs to the NMJs. Aldicarb, an inhibitor of acetylcholinesterase, causes paralysis in C. elegans due to accumulation of acetylcholine (ACh) at the locomotory NMJs, but exogenous serotonin inhibits paralysis induced by aldicarb. By contrast, exogenous serotonin does not reduce the paralysis induced by levamisole, a specific agonist of the nicotinic ACh receptor UNC-29 in the bodywall muscles. This suggests that serotonin signaling inhibits ACh release by the motor neurons. Studies showed that the inhibitory and stimulatory serotonin signals to NMJ neurotransmission arise from distinct serotonergic neurons; endogenous serotonin released from the ADF neurons stimulates ACh synaptic transmission at the NMJs, whereas serotonin from the NSM/RIH/AIM neurons inhibits the stimulatory serotonin inputs (Govorunova et al., 2010).
8
ADLL
Sensory neuron
Lateral ganglia in headAmphid neurons, dual (AdL) ciliated sensory endings. Project directly to ring, take up FITC.- FLP-4; FMRFamide-like neuropeptide
- FLP-21; FMRFamide-like neuropeptide
- NLP-7; neuropeptide-like peptide
- NLP-8; neuropeptide-like peptide
- NLP-10; neuropeptide-like peptide
(Li and Kim, 2008; Rogers et al., 2003; Nathoo et al., 2001)
- INX-18
- INX-19
- UNC-9
(Altun et al., 2009; Chuang et al., 2007; Liu et al., 2006)
- GCY-21; transmembrane receptor guanylate cyclase
- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)-cation selective
- OCR-1; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- OCR-2; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- SRB-6; G protein-coupled seven transmembrane receptor
- SRE-1; G protein-coupled seven transmembrane receptor
- SRH-132; G protein-coupled seven transmembrane receptor
- SRH-220; G protein-coupled seven transmembrane receptor
- SRI-51; G protein-coupled seven transmembrane receptor
- SRO-1; G protein-coupled seven transmembrane receptor
(Wormbase; Tobin et al., 2002; de Bono et al., 2002; Colbert et al., 1997; Troemel et al., 1995)
- ASH, ADL, ASK and ASE sensory neurons are responsible for the detection of certain chemical repellents. ASH plays a major role in this avoidance, whereas ADL, ASK and ASE play minor roles that are only evident when ASH is missing (de Bono and Maricq, 2005; Hiliard et al., 2002; Sambongi et al., 1999; Bargmann et al., 1990). ADL mediate avoidance behavior from heavy metals (Cd++ and Cu++) (Sambongi et al., 1999), odors (e.g. octanol) (Troemel et al., 1995), high osmotic strength and SDS.
- ASH and ADL are proposed to mediate social feeding behavior in response to repulsive cues (e.g. high O2 levels); ablation of ASH and ADL abolishes social feeding behavior transforming social animals to solitary feeders; avoidance of high O2 levels that facilitate aggregation is promoted by OCR-2 and OSM-9, and the transmembrane protein ODR-4, acting in the nociceptive neurons ASH and ADL. ASH and ADL, in turn, transmit information about aversive stimuli in the environment to a circuit that is responsible for aggregation, rapid locomotion, and food bordering behavior (Rogers et al., 2006; de Bono et al., 2002).
- Balances between at least four sensory cell types, ASE, ASI, ASH, ADF and perhaps ADL, modulate the response to NaCl (Hukema et al, 2006).
9
ADLR
Sensory neuron
Lateral ganglia in headAmphid neurons, dual (AdL) ciliated sensory endings. Project directly to ring, take up FITC.- FLP-4; FMRFamide-like neuropeptide
- FLP-21; FMRFamide-like neuropeptide
- NLP-7; neuropeptide-like peptide
- NLP-8; neuropeptide-like peptide
- NLP-10; neuropeptide-like peptide
(Li and Kim, 2008; Rogers et al., 2003; Nathoo et al., 2001)
- INX-18
- INX-19
- UNC-9
(Altun et al., 2009; Chuang et al., 2007; Liu et al., 2006)
- GCY-21; transmembrane receptor guanylate cyclase
- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)-cation selective
- OCR-1; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- OCR-2; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)
- SRB-6; G protein-coupled seven transmembrane receptor
- SRE-1; G protein-coupled seven transmembrane receptor
- SRH-132; G protein-coupled seven transmembrane receptor
- SRH-220; G protein-coupled seven transmembrane receptor
- SRI-51; G protein-coupled seven transmembrane receptor
- SRO-1; G protein-coupled seven transmembrane receptor
(Wormbase; Tobin et al., 2002; de Bono et al., 2002; Colbert et al., 1997; Troemel et al., 1995)
- ASH, ADL, ASK and ASE sensory neurons are responsible for the detection of certain chemical repellents. ASH plays a major role in this avoidance, whereas ADL, ASK and ASE play minor roles that are only evident when ASH is missing (de Bono and Maricq, 2005; Hiliard et al., 2002; Sambongi et al., 1999; Bargmann et al., 1990). ADL mediate avoidance behavior from heavy metals (Cd++ and Cu++) (Sambongi et al., 1999), odors (e.g. octanol) (Troemel et al., 1995), high osmotic strength and SDS.
- ASH and ADL are proposed to mediate social feeding behavior in response to repulsive cues (e.g. high O2 levels); ablation of ASH and ADL abolishes social feeding behavior transforming social animals to solitary feeders; avoidance of high O2 levels that facilitate aggregation is promoted by OCR-2 and OSM-9, and the transmembrane protein ODR-4, acting in the nociceptive neurons ASH and ADL. ASH and ADL, in turn, transmit information about aversive stimuli in the environment to a circuit that is responsible for aggregation, rapid locomotion, and food bordering behavior (Rogers et al., 2006; de Bono et al., 2002).
- Balances between at least four sensory cell types, ASE, ASI, ASH, ADF and perhaps ADL, modulate the response to NaCl (Hukema et al, 2006).
10
AFDL
Sensory neuron
Lateral ganglia in headAmphid neurons with finger-like (AfD) ciliated endings embedded in the amphid sheath cell.- Glutamate
- FLP-6; FMRFamide-like neuropeptide
- NLP-7; neuropeptide-like peptide
- NLP-21; neuropeptide-like peptide
(Ohnishi et al, 2011; Li and Kim, 2008; Rogers et al., 2003; Nathoo et al., 2001)
- INX-19
(Chuang et al., 2007)
- GCY-8; transmembrane receptor guanylate cyclase
- GCY-18; transmembrane receptor guanylate cyclase
- GCY-23; transmembrane receptor guanylate cyclase
- GCY-29; transmembrane receptor guanylate cyclase
(Wormbase; Ortiz et al., 2006; Inada et al., 2006; Yu et al., 1997)
- Functions in thermotaxis: AFD neurons are the main thermosensors in C. elegans and laser ablation of the AFD pair makes most animals athermotactic (Ma and Shen, 2012; Beverly et al, 2011; Kuhara et al, 2008; Satterlee et al., 2004; Mori and Ohshima, 1995). After cultivation at a uniform temperature (Tc) with sufficient food, animals preferentially migrate to their cultivation temperature (Tc) when placed on a thermal gradient, and move isothermally at this temperature (Hedgecock and Russell 1975). Animals sense and record their Tc by AFD (major thermosensory), AWC and ASI neurons (Beverly et al, 2011; Kuhara et al., 2008; Biron et al., 2008). This memory is plastic and can be reset upon cultivation at a different temperature (Hedgecock and Russell 1975). The AFD, AWC and ASI neurons may to act in concert to increase turning rate when animals encounter higher temperatures than Tc on a gradient to move down the gradient toward colder temperatures near Tc (negative thermotaxis) (Biron et al., 2008). To track isotherms, animals do not actively pursue isothermal alignment, but once serendipitously aligned along an isotherm (at T=Tc), they track by suppressing turns (Luo et al., 2006).This may be achieved by the animal increasing its reversal frequency when it detects a rise in temperature, which activates Ca++ influx in AFD via cGMP-dependent TAX-2/TAX-4 cation channel (cGMP production in AFD is redundantly carried out by GCY-8, GCY-18 and GCY-23 and gcy-23, gcy-8, and gcy-18 triple mutants show a cryophilic or athermotactic phenotype) (Inada et al., 2006). AFD neurons respond to thermal stimuli above Tc with continuous, graded calcium signals in a deterministic and highly reproducible way. The thermal information is transmitted by AFD and AWC to AIY interneurons for information processing, and AIY neurons, in turn, transmit it to AIZ and RIA interneurons for further processing. It is suggested that activation of AFD neurons by warming above Tc induces the animal to reverse by inhibiting AIY since loss of AFD (by ablation) suppresses spontaneous reversals in an AIY-dependent manner, while loss of AIY (by ablation) increases spontaneous reversals (de Bono and Maricq 2005; Tsalik and Hobert, 2003).
- Functions in locomotion: Laser ablations of AFD cause hyporeversal phenotype (Tsalik and Hobert, 2003).
- AFD, BAG and ASE are primary CO2 sensors: AFD and BAG neurons together stimulate turning when CO2 rises and inhibit turning when CO2 falls (Bretscher et al., 2011).
- May be involved in social feeding since disruption of tax-2 expression in AFD, AQR, ASE and BAG neurons disrupts social feeding (Coates and de Bono, 2002).
11
AFDR
Sensory neuron
Lateral ganglia in headAmphid neurons with finger-like (AfD) ciliated endings embedded in the amphid sheath cell.- Glutamate
- FLP-6; FMRFamide-like neuropeptide
- NLP-7; neuropeptide-like peptide
- NLP-21; neuropeptide-like peptide
(Ohnishi et al, 2011; Li and Kim, 2008; Rogers et al., 2003; Nathoo et al., 2001)
- INX-19
(Chuang et al., 2007)
- GCY-8; transmembrane receptor guanylate cyclase
- GCY-18; transmembrane receptor guanylate cyclase
- GCY-23; transmembrane receptor guanylate cyclase
- GCY-29; transmembrane receptor guanylate cyclase
(Wormbase; Ortiz et al., 2006; Inada et al., 2006; Yu et al., 1997)
- Functions in thermotaxis: AFD neurons are the main thermosensors in C. elegans and laser ablation of the AFD pair makes most animals athermotactic (Ma and Shen, 2012; Beverly et al, 2011; Kuhara et al, 2008; Satterlee et al., 2004; Mori and Ohshima, 1995). After cultivation at a uniform temperature (Tc) with sufficient food, animals preferentially migrate to their cultivation temperature (Tc) when placed on a thermal gradient, and move isothermally at this temperature (Hedgecock and Russell 1975). Animals sense and record their Tc by AFD (major thermosensory), AWC and ASI neurons (Beverly et al, 2011; Kuhara et al., 2008; Biron et al., 2008). This memory is plastic and can be reset upon cultivation at a different temperature (Hedgecock and Russell 1975). The AFD, AWC and ASI neurons may to act in concert to increase turning rate when animals encounter higher temperatures than Tc on a gradient to move down the gradient toward colder temperatures near Tc (negative thermotaxis) (Biron et al., 2008). To track isotherms, animals do not actively pursue isothermal alignment, but once serendipitously aligned along an isotherm (at T=Tc), they track by suppressing turns (Luo et al., 2006).This may be achieved by the animal increasing its reversal frequency when it detects a rise in temperature, which activates Ca++ influx in AFD via cGMP-dependent TAX-2/TAX-4 cation channel (cGMP production in AFD is redundantly carried out by GCY-8, GCY-18 and GCY-23 and gcy-23, gcy-8, and gcy-18 triple mutants show a cryophilic or athermotactic phenotype) (Inada et al., 2006). AFD neurons respond to thermal stimuli above Tc with continuous, graded calcium signals in a deterministic and highly reproducible way. The thermal information is transmitted by AFD and AWC to AIY interneurons for information processing, and AIY neurons, in turn, transmit it to AIZ and RIA interneurons for further processing. It is suggested that activation of AFD neurons by warming above Tc induces the animal to reverse by inhibiting AIY since loss of AFD (by ablation) suppresses spontaneous reversals in an AIY-dependent manner, while loss of AIY (by ablation) increases spontaneous reversals (de Bono and Maricq 2005; Tsalik and Hobert, 2003).
- Functions in locomotion: Laser ablations of AFD cause hyporeversal phenotype (Tsalik and Hobert, 2003).
- AFD, BAG and ASE are primary CO2 sensors: AFD and BAG neurons together stimulate turning when CO2 rises and inhibit turning when CO2 falls (Bretscher et al., 2011).
- May be involved in social feeding since disruption of tax-2 expression in AFD, AQR, ASE and BAG neurons disrupts social feeding (Coates and de Bono, 2002).
12
AIAL
Interneuron
Ventral ganglion in headRing interneurons. AIAL and AIAR are located anteriorly to the excretory cell, at the ventral side of the terminal bulb of pharynx. They are seen very close to each other on the right and left sides of the A-P midline in lateral view.- Acetylcholine
- FLP-1; FMRFamide-like neuropeptide
- FLP-2; FMRFamide-like neuropeptide
- INS-1; insulin-like peptide. Important in the neuropeptide feedback loop between AWC and AIA.
(Chalasani et al., 2010; Li and Kim, 2008; Altun-Gultekin et al., 2001; Nelson et al., 1998)
None yet reported, although described to have gap junctions in adult animals (MoW)- GLR-2; AMPA-type ionotropic (Na+/K+) glutamate receptor subunit
- GLC-3; L-glutamate-gated chloride channel subunit (ionotropic glutamate rec subunit). AWC-AIA synapse is an inhibitory, glutamatergic synapse via GLC-3, whose function is acutely modulated by NLP-1/NPR-11 signaling
- GCY-28; transmembrane receptor guanylate cyclase
- MGL-1; group II metabotropic glutamate receptor
- NPR-5; receptor for FLP-18 peptides
- NPR-11; candidate receptor for NLP-1
- SCD-2; ALK receptor tyrosine kinase
- SRA-11; G protein-coupled seven transmembrane receptor
(Wormbase; Shinkai et al, 2011; Chalasani et al., 2010; Glauser and Goodman, 2010; Greer et al, 2008; Altun-Gultekin et al., 2001; Brockie et al., 2001)
Integration of information from amphid sensory neurons; AIAL/R is one of the four first layer amphid interneuron pairs (AIA, AIB, AIY, and AIZ) that receives and processes synaptic output from the amphid sensory neurons towards a behavioral response. AIA pair is suggested to sum inputs from various chemosensory neurons before passing the information on to AIB pair, which synapses onto motor neurons (AIA-AIB connections are likely to be inhibitory) (Wakayabashi et al., 2004). AIA is the main target of ASI outputs.
- AIA interneurons integrate multiple sensory cues to adjust behavioral choices and play an important role in learning (e.g. salt chemotaxis learning) and behavioral plasticity (Shinkai et al, 2011; Tomioka et al., 2006).
- Functions in locomotion: in isothermal tracking, AIB and AIZ interneurons promote turns while AIY and AIA interneurons inhibit turns (Garrity et al., 2010).
- The AIA neurons are required for chemotaxis to indole ascarosides, downstream of the ASK sensory neuron (Macosko et al., 2009).
13
AIAR
Interneuron
Ventral ganglion in headRing interneurons. AIAL and AIAR are located anteriorly to the excretory cell, at the ventral side of the terminal bulb of pharynx. They are seen very close to each other on the right and left sides of the A-P midline in lateral view.- Acetylcholine
- FLP-1; FMRFamide-like neuropeptide
- FLP-2; FMRFamide-like neuropeptide
- INS-1; insulin-like peptide. Important in the neuropeptide feedback loop between AWC and AIA.
(Chalasani et al., 2010; Li and Kim, 2008; Altun-Gultekin et al., 2001; Nelson et al., 1998)
None yet reported, although described to have gap junctions in adult animals (MoW)- GLR-2; AMPA-type ionotropic (Na+/K+) glutamate receptor subunit
- GLC-3; L-glutamate-gated chloride channel subunit (ionotropic glutamate rec subunit). AWC-AIA synapse is an inhibitory, glutamatergic synapse via GLC-3, whose function is acutely modulated by NLP-1/NPR-11 signaling
- GCY-28; transmembrane receptor guanylate cyclase
- MGL-1; group II metabotropic glutamate receptor
- NPR-5; receptor for FLP-18 peptides
- NPR-11; candidate receptor for NLP-1
- SCD-2; ALK receptor tyrosine kinase
- SRA-11; G protein-coupled seven transmembrane receptor
(Wormbase; Shinkai et al, 2011; Chalasani et al., 2010; Glauser and Goodman, 2010; Greer et al, 2008; Altun-Gultekin et al., 2001; Brockie et al., 2001)
Integration of information from amphid sensory neurons; AIAL/R is one of the four first layer amphid interneuron pairs (AIA, AIB, AIY, and AIZ) that receives and processes synaptic output from the amphid sensory neurons towards a behavioral response. AIA pair is suggested to sum inputs from various chemosensory neurons before passing the information on to AIB pair, which synapses onto motor neurons (AIA-AIB connections are likely to be inhibitory) (Wakayabashi et al., 2004). AIA is the main target of ASI outputs.
- AIA interneurons integrate multiple sensory cues to adjust behavioral choices and play an important role in learning (e.g. salt chemotaxis learning) and behavioral plasticity (Shinkai et al, 2011; Tomioka et al., 2006).
- Functions in locomotion: in isothermal tracking, AIB and AIZ interneurons promote turns while AIY and AIA interneurons inhibit turns (Garrity et al., 2010).
- The AIA neurons are required for chemotaxis to indole ascarosides, downstream of the ASK sensory neuron (Macosko et al., 2009).
14
AIBL
Interneuron
Lateral ganglia in headAmphid interneuron- FLP-20; FMRFamide-like neuropeptide
(Li and Kim, 2008)
- INX-1
(Altun et al, 2009)
- GLR-1; glutamate-activated cation channel
- GLR-2; glutamate-activated cation channel
- Possibly GLR-5; kainate (non-NMDA)-type ionotropic glutamate receptor
- Possibly GGR-1; GABA-A/glycine receptor-like protein
- Possibly MGL-2; metabotropic glutamate receptor
(Fujiwara et al., 1996; Maricq et al., 1995; Brockie et al., 2001; Kang and Avery, 2009)
Integration of information from amphid sensory neurons; AIBL/R is one of the four first layer amphid interneuron pairs (AIA, AIB, AIY, and AIZ) that receives and processes synaptic output from the amphid sensory neurons. AIA pair is suggested to sum inputs from various chemosensory neurons before passing the information on to AIB pair, which synapses onto motor neurons (AIA-AIB connections are likely to be inhibitory) (Wakayabashi et al., 2004).
- Functions in locomotion: Among the interneurons, AIA and AIY inhibit turns, AIB and AIZ promote turns, and ablation of RIA does not affect turning rates (Garrity et al., 2010; Wakayabashi et al., 2004).
- Functions in information processing within the AWC-AIB-AIY circuit that controls food- and odor-evoked behaviors (Chalasani et al., 2007). After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later, as reversals and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005). AWC neurons are activated by odor removal and activate the AIB interneurons through AMPA-type glutamate receptors, while they inhibit AIY interneurons through glutamate-gated chloride channels; odor presentation relieves this inhibition and results in activation of AIY interneurons. The opposite regulation of AIY and AIB interneurons generates a coordinated behavioral response.
- Regulates lifespan and starvation response (starvation induces heat-shock resistance, oxidative stress resistance and extension of life-span). Certain amino acids (such as leu, gln, ala, val, ile) might act as signaling molecules that modulate the starvation response by acting as anti-hunger signals. The amino acid signal activates AIY neurons and inhibits AIB neurons by modulating the activities of MGL-1 and MGL-2 metabotropic glutamate receptors, respectively. AIY then inhibits the starvation response, whereas AIB activates it (Shen et al, 2010; Kang and Avery, 2009).
15
AIBR
Interneuron
Lateral ganglia in headAmphid interneuron- FLP-20; FMRFamide-like neuropeptide
(Li and Kim, 2008)
- INX-1
(Altun et al, 2009)
- GLR-1; glutamate-activated cation channel
- GLR-2; glutamate-activated cation channel
- Possibly GLR-5; kainate (non-NMDA)-type ionotropic glutamate receptor
- Possibly GGR-1; GABA-A/glycine receptor-like protein
- Possibly MGL-2; metabotropic glutamate receptor
(Fujiwara et al., 1996; Maricq et al., 1995; Brockie et al., 2001; Kang and Avery, 2009)
Integration of information from amphid sensory neurons; AIBL/R is one of the four first layer amphid interneuron pairs (AIA, AIB, AIY, and AIZ) that receives and processes synaptic output from the amphid sensory neurons. AIA pair is suggested to sum inputs from various chemosensory neurons before passing the information on to AIB pair, which synapses onto motor neurons (AIA-AIB connections are likely to be inhibitory) (Wakayabashi et al., 2004).
- Functions in locomotion: Among the interneurons, AIA and AIY inhibit turns, AIB and AIZ promote turns, and ablation of RIA does not affect turning rates (Garrity et al., 2010; Wakayabashi et al., 2004).
- Functions in information processing within the AWC-AIB-AIY circuit that controls food- and odor-evoked behaviors (Chalasani et al., 2007). After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later, as reversals and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005). AWC neurons are activated by odor removal and activate the AIB interneurons through AMPA-type glutamate receptors, while they inhibit AIY interneurons through glutamate-gated chloride channels; odor presentation relieves this inhibition and results in activation of AIY interneurons. The opposite regulation of AIY and AIB interneurons generates a coordinated behavioral response.
- Regulates lifespan and starvation response (starvation induces heat-shock resistance, oxidative stress resistance and extension of life-span). Certain amino acids (such as leu, gln, ala, val, ile) might act as signaling molecules that modulate the starvation response by acting as anti-hunger signals. The amino acid signal activates AIY neurons and inhibits AIB neurons by modulating the activities of MGL-1 and MGL-2 metabotropic glutamate receptors, respectively. AIY then inhibits the starvation response, whereas AIB activates it (Shen et al, 2010; Kang and Avery, 2009).
16
AIML
Interneuron
Ventral ganglion in headRing interneurons bridging between sensory inputs and command outputs. Excessive neurites that connect AIML and AIMR early in development are pruned during larval stages (along with RIF, they are the only other neuron pair that is subject to developmental neurite pruning in C. elegans) (Hayashi et al., 2009; Kage et al., 2005).- Serotonin
- FLP-10; FMRFamide-like peptide
- FLP-22; FMRFamide-like peptide
- INS-1; insulin-like peptide
(Loer, 2010; Li and Kim, 2008; Sze et al., 2000; Duerr et al., 1999)
- INX-19
(Chuang et al, 2007)
- GCY-18; guanylate cyclase
(Wormbase; Ortiz et al., 2006)
Serotonin secreted by both synaptic vesicles and dense core vesicles seem to diffuse readily to the extrasynaptic space adjacent to the AIM and RIH neurons. Among 5 classes (NSM, ADF, HSN, AIM, RIH) of serotonergic neurons, RIH and AIM mainly take up serotonin from outside -produced by the remaining serotonergic neurons- via the membrane serotonin transporter (MOD-5/SERT), rather than synthesizing it themselves (Jafari et al., 2011).
- RIH and AIM possibly function as temporal–spatial regulators of extrasynaptic serotonin to modulate the activity of the behavioral circuits. In Caenorhabditis elegans, serotonin signaling is required for coupling feeding and the memory of prior food availability to the rate of locomotion. Following food deprivation, wild-type animals show markedly decreased locomotion on their return to a lawn of bacterial food. While serotonin-deficient mutants exhibit less slowing response (become insensitive), mutants of mod-5/SERT exaggerate the slowing response (become hypersensitive). Transgenic expression of mod-5/SERT in the serotonin-absorbing neurons fully corrects this exaggerated behavior, suggesting AIM and RIH regulate the availability of serotonin for neurotransmission (Jafari et al., 2011).
- Functions in swim initiation; Behaviorally, ablation of VC-4, VC-5, AIM, or NSM neurons delays swim onset, implicating serotonin signaling in this behavior (Vidal-Gadea et al., 2011).
17
AIMR
Interneuron
Ventral ganglion in headRing interneurons bridging between sensory inputs and command outputs. Excessive neurites that connect AIML and AIMR early in development are pruned during larval stages (along with RIF, they are the only other neuron pair that is subject to developmental neurite pruning in C. elegans) (Hayashi et al., 2009; Kage et al., 2005).- Serotonin
- FLP-10; FMRFamide-like peptide
- FLP-22; FMRFamide-like peptide
- INS-1; insulin-like peptide
(Loer, 2010; Li and Kim, 2008; Sze et al., 2000; Duerr et al., 1999)
- INX-19
(Chuang et al, 2007)
- GCY-18; guanylate cyclase
(Wormbase; Ortiz et al., 2006)
Serotonin secreted by both synaptic vesicles and dense core vesicles seem to diffuse readily to the extrasynaptic space adjacent to the AIM and RIH neurons. Among 5 classes (NSM, ADF, HSN, AIM, RIH) of serotonergic neurons, RIH and AIM mainly take up serotonin from outside -produced by the remaining serotonergic neurons- via the membrane serotonin transporter (MOD-5/SERT), rather than synthesizing it themselves (Jafari et al., 2011).
- RIH and AIM possibly function as temporal–spatial regulators of extrasynaptic serotonin to modulate the activity of the behavioral circuits. In Caenorhabditis elegans, serotonin signaling is required for coupling feeding and the memory of prior food availability to the rate of locomotion. Following food deprivation, wild-type animals show markedly decreased locomotion on their return to a lawn of bacterial food. While serotonin-deficient mutants exhibit less slowing response (become insensitive), mutants of mod-5/SERT exaggerate the slowing response (become hypersensitive). Transgenic expression of mod-5/SERT in the serotonin-absorbing neurons fully corrects this exaggerated behavior, suggesting AIM and RIH regulate the availability of serotonin for neurotransmission (Jafari et al., 2011).
- Functions in swim initiation; Behaviorally, ablation of VC-4, VC-5, AIM, or NSM neurons delays swim onset, implicating serotonin signaling in this behavior (Vidal-Gadea et al., 2011).
18
AINL
Interneuron
Lateral ganglia in headRing interneuron- Possibly glutamate (the neuron may be AVJ instead)
- FLP-19; FMRFamide-like peptide
(Loer, 2010; Li and Kim, 2008; Lee et al., 1999)
- INX-4
- INX-8
- INX-17
- UNC-7
- UNC-9
(Altun et al., 2009)
UnknownAIN is presynaptic to AFD, however, the significance of this connection is not yet clear.
19
AINR
Interneuron
Lateral ganglia in headRing interneuron- Possibly glutamate (the neuron may be AVJ instead)
- FLP-19; FMRFamide-like peptide
(Loer, 2010; Li and Kim, 2008; Lee et al., 1999)
- INX-4
- INX-8
- INX-17
- UNC-7
- UNC-9
(Altun et al., 2009)
UnknownAIN is presynaptic to AFD, however, the significance of this connection is not yet clear.
20
AIYL
Interneuron
Ventral ganglion in headAmphid interneuron- Acetylcholine
- FLP-1; FMRFamide-like peptide
- FLP-18; FMRFamide-like peptide
(Li and Kim, 2008; Rogers et al., 2003; Altun-Gultekin et al., 2001)
- INX-1
- INX-7
- INX-19
- UNC-9
(Altun et al., 2009; Chuang et al., 2007)
- ACR-14; nicotinic AChR alpha subunit
- C50F7.1; tachykinin receptor-like
- CKR-2; cholecystokinin receptor-like
- GAR-2; muscarinic AChR
- GCY-1; transmembrane receptor guanylate cyclase
- GLC-3; L-glutamate-gated chloride channel subunit (ionotropic glutamate rec subunit) - LAT-1; predicted latrophilin
- LGC-38; GABA-gated chloride channel
- MGL-1; metabotropic glutamate receptor
- MOD-1; serotonin receptor (serotonin-gated Cl- channel)
- NPR-11; candidate receptor for NLP-1
- NPR-14; tachykinin receptor-like
- SER-2; tyramine receptor (expresses two separate splice variants)
- SRA-11; G protein-coupled seven transmembrane receptor
(Chalasani et al., 2010; Ortiz et al., 2006; Wenick and Hobert, 2004; Tsalik et al., 2003; Altun-Gultekin et al., 2001; Troemel et al., 1995)
AIY neurons are postsynaptic to several olfactory and gustatory sensory neurons, such as ASE, ASI, AWA, AWC and AFD, and are presumed to play an integrative role, processing multiple streams of sensory information. They have a key role in behavioral plasticity in paradigms in which starvation is paired with a thermal or chemical cue (de Bono and Maricq, 2005; Ishihara et al., 2002).
- Functions in thermotaxis; laser ablation of the AIY pair makes the animals cryophilic and impaired in isothermal tracking (Mori and Ohshima, 1995). The thermal information sensed by AFD, AWC and ASI is processed in downstream interneurons AIY, AIZ, RIA, and also likely in AIA and AIB (the first three are considered as thermotaxis core interneurons) (Ma and Shen, 2012; Sasakura and Mori, 2012; Ardiel and Rankin, 2011; Beverly et al, 2011). The activation of AIY neuron-signaling drive warm-seeking (thermophilic) movement, the activation of AIZ neuron-signaling drive cold-seeking (cryophilic) movement, while the counterbalancing between the AIY and AIZ signals through the RIA interneuron leads to execution of motor output in response to thermal signals (Ohnishi et al, 2011; Kuhara et al, 2008). AIY is the major postsynaptic partner of AWC and AFD. AWC-derived glutamatergic signals stimulate AIY and induce migration to warmer temperature (Ohnishi et al, 2011). AFD transmits both inhibitory and stimulatory signals to AIY such that calcium concentration threshold in AFD acts as a switch for opposing neural signals to AIY that direct opposite (warm- or cold-seeking) behaviors (Kuhara et al., 2011). AFD-derived glutamatergic signals inhibit AIY through activation of GLC-3 glutamate-gated chloride channel and drive migration towards colder temperature. Recent studies suggest this AFD / AIY synapse may have evolved for reliable transmission of a scaled-down temperature signal from AFD, enabling AIY to monitor and integrate temperature with other sensory inputs (Narayan et al, 2011). The molecular mechanism of the stimulatory signaling between AFD and AIY is currently unknown.
- Functions in locomotion: Suppresses turns and reversals enhancing smooth forward movements and dispersal, since laser ablation of AIY increases reversals and turns, disrupts dispersal behavior and shortens forward movements (Gray et al., 2005; Wakayabashi et al., 2004 ; Tsalik and Hobert, 2003). Similar to AIY, laser ablations of ASE or AIA increases the frequency of reversals while ablations of AWC, AIB or AIZ decreases the frequency of reversals (ablations of RIA does not affect turning rates). Since ablations of AIY and AIZ (the major postsynaptic partner of AIY) cause opposite behavioral phenotpes, the synapses made by AIY onto AIZ may be inhibitory. Alternatively, signals fom AIY to AIZ are processed in a downstream neuron such as RIA (de Bono and Maricq 2005).
- Functions in information processing within the AWC-AIB-AIY circuit that controls food- and odor-evoked behaviors (Chalasani et al., 2007). AWC neurons are activated by food or odor removal and activate the AIB interneurons through AMPA-type glutamate receptors, while they inhibit AIY interneurons through glutamate-gated chloride channels; odor presentation relieves this inhibition and results in activation of AIY interneurons. The opposite regulation of AIY and AIB interneurons generates a coordinated behavioral response.
- Regulates lifespan and starvation response (starvation induces heat-shock resistance, oxidative stress resistance and extension of life-span). Certain amino acids (such as leu, gln, ala, val, ile) might act as signaling molecules that modulate the starvation response by acting as anti-hunger signals. The amino acid signal activates AIY neurons and inhibits AIB neurons by modulating the activities of MGL-1 and MGL-2 metabotropic glutamate receptors in these neurons, respectively. AIY then inhibits the starvation response, whereas AIB activates it (Shen et al, 2010; Kang and Avery, 2009).
21
AIYR
Interneuron
Ventral ganglion in headAmphid interneuron- Acetylcholine
- FLP-1; FMRFamide-like peptide
- FLP-18; FMRFamide-like peptide
(Li and Kim, 2008; Rogers et al., 2003; Altun-Gultekin et al., 2001)
- INX-1
- INX-7
- INX-19
- UNC-9
(Altun et al., 2009; Chuang et al., 2007)
- ACR-14; nicotinic AChR alpha subunit
- C50F7.1; tachykinin receptor-like
- CKR-2; cholecystokinin receptor-like
- GAR-2; muscarinic AChR
- GCY-1; transmembrane receptor guanylate cyclase
- GLC-3; L-glutamate-gated chloride channel subunit (ionotropic glutamate rec subunit) - LAT-1; predicted latrophilin
- LGC-38; GABA-gated chloride channel
- MGL-1; metabotropic glutamate receptor
- MOD-1; serotonin receptor (serotonin-gated Cl- channel)
- NPR-11; candidate receptor for NLP-1
- NPR-14; tachykinin receptor-like
- SER-2; tyramine receptor (expresses two separate splice variants)
- SRA-11; G protein-coupled seven transmembrane receptor
(Chalasani et al., 2010; Ortiz et al., 2006; Wenick and Hobert, 2004; Tsalik et al., 2003; Altun-Gultekin et al., 2001; Troemel et al., 1995)
AIY neurons are postsynaptic to several olfactory and gustatory sensory neurons, such as ASE, ASI, AWA, AWC and AFD, and are presumed to play an integrative role, processing multiple streams of sensory information. They have a key role in behavioral plasticity in paradigms in which starvation is paired with a thermal or chemical cue (de Bono and Maricq, 2005; Ishihara et al., 2002).
- Functions in thermotaxis; laser ablation of the AIY pair makes the animals cryophilic and impaired in isothermal tracking (Mori and Ohshima, 1995). The thermal information sensed by AFD, AWC and ASI is processed in downstream interneurons AIY, AIZ, RIA, and also likely in AIA and AIB (the first three are considered as thermotaxis core interneurons) (Ma and Shen, 2012; Sasakura and Mori, 2012; Ardiel and Rankin, 2011; Beverly et al, 2011). The activation of AIY neuron-signaling drive warm-seeking (thermophilic) movement, the activation of AIZ neuron-signaling drive cold-seeking (cryophilic) movement, while the counterbalancing between the AIY and AIZ signals through the RIA interneuron leads to execution of motor output in response to thermal signals (Ohnishi et al, 2011; Kuhara et al, 2008). AIY is the major postsynaptic partner of AWC and AFD. AWC-derived glutamatergic signals stimulate AIY and induce migration to warmer temperature (Ohnishi et al, 2011). AFD transmits both inhibitory and stimulatory signals to AIY such that calcium concentration threshold in AFD acts as a switch for opposing neural signals to AIY that direct opposite (warm- or cold-seeking) behaviors (Kuhara et al., 2011). AFD-derived glutamatergic signals inhibit AIY through activation of GLC-3 glutamate-gated chloride channel and drive migration towards colder temperature. Recent studies suggest this AFD / AIY synapse may have evolved for reliable transmission of a scaled-down temperature signal from AFD, enabling AIY to monitor and integrate temperature with other sensory inputs (Narayan et al, 2011). The molecular mechanism of the stimulatory signaling between AFD and AIY is currently unknown.
- Functions in locomotion: Suppresses turns and reversals enhancing smooth forward movements and dispersal, since laser ablation of AIY increases reversals and turns, disrupts dispersal behavior and shortens forward movements (Gray et al., 2005; Wakayabashi et al., 2004 ; Tsalik and Hobert, 2003). Similar to AIY, laser ablations of ASE or AIA increases the frequency of reversals while ablations of AWC, AIB or AIZ decreases the frequency of reversals (ablations of RIA does not affect turning rates). Since ablations of AIY and AIZ (the major postsynaptic partner of AIY) cause opposite behavioral phenotpes, the synapses made by AIY onto AIZ may be inhibitory. Alternatively, signals fom AIY to AIZ are processed in a downstream neuron such as RIA (de Bono and Maricq 2005).
- Functions in information processing within the AWC-AIB-AIY circuit that controls food- and odor-evoked behaviors (Chalasani et al., 2007). AWC neurons are activated by food or odor removal and activate the AIB interneurons through AMPA-type glutamate receptors, while they inhibit AIY interneurons through glutamate-gated chloride channels; odor presentation relieves this inhibition and results in activation of AIY interneurons. The opposite regulation of AIY and AIB interneurons generates a coordinated behavioral response.
- Regulates lifespan and starvation response (starvation induces heat-shock resistance, oxidative stress resistance and extension of life-span). Certain amino acids (such as leu, gln, ala, val, ile) might act as signaling molecules that modulate the starvation response by acting as anti-hunger signals. The amino acid signal activates AIY neurons and inhibits AIB neurons by modulating the activities of MGL-1 and MGL-2 metabotropic glutamate receptors in these neurons, respectively. AIY then inhibits the starvation response, whereas AIB activates it (Shen et al, 2010; Kang and Avery, 2009).
22
AIZL
Interneuron
Ventral ganglion in headRing Interneurons- SER-2; tyramine receptor (expresses two separate splice variants)
(Tsalik et al., 2003)
Integration of information from amphid sensory neurons; AIZL/R is one of the four first layer amphid interneuron pairs (AIA, AIB, AIY, and AIZ) that receive and process synaptic output from the amphid sensory neurons towards a behavioral response. Along with the second layer RIA and RIB interneurons, AIZ also receives heavy input from the first layer AIA, AIB, AIY interneurons and itself sends inputs to RIA and RIB. Additionally, it synapses directly onto SMB head motor neurons .
- Functions in thermotaxis; laser ablation of the AIZ pair makes the animals thermophilic (Mori and Ohshima, 1995). The thermal information sensed by AFD, AWC and ASI is processed in downstream interneurons AIY, AIZ, RIA, and also likely in AIA and AIB (the first three are considered as thermotaxis core interneurons) (Ma and Shen, 2012; Sasakura and Mori, 2012; Ardiel and Rankin, 2011; Beverly et al, 2011). In the original two-drive model of thermotaxis, activation of AIY neuron-signaling drive warm-seeking (thermophilic) movement, activation of AIZ neuron-signaling drive cold-seeking (cryophilic) movement, while the counterbalancing between the AIY and AIZ signals possibly through the RIA interneuron leads to execution of motor output in response to thermal signals (Ohnishi et al, 2011; Kuhara et al, 2008). However, there is evidence that a thermophilic drive at temperatures below cultivation temperature (Tc) does not exist and regulation of thermotaxis involves a more complex circuit (Ma and Shen, 2012; Ryu and Samuel, 2002). Around Tc animals track isothermally on a gradient and above Tc they display negative thermotaxis by regulating their turning frequency and run duration. It has been suggested that thermosensory signals from AFD, AWC and ASI converge on the AIB interneuron in negative thermotaxis, whereas the AIY/AIZ/RIA network may function mainly in isothermal tracking (Ma and Shen, 2012).
During associative learning between temperature and feeding state, starvation, a conditioning factor, downregulates AIZ activity through calcineurin/TAX-6, a calcium-activated phosphatase, leading to avoidance from Tc (Kuhara and Mori, 2006).
- Functions in locomotion: Laser ablations of AIZ, like AIB and AWC, cause decrease in reversals and lead to large increases in the duration of forward movement (Tsalik and Hobert, 2003; Iino and Yoshida, 2009). Since ablations of AIY and AIZ cause opposite behavioral phenotpes, the synapses made by AIY onto AIZ may be inhibitory. Alternatively, signals fom AIY to AIZ could be processed in a downstream neuron such as RIA (de Bono and Maricq 2005).
- Functions in chemotaxis: Laser ablations of AIZ cause defects in the weathervane (slow turning of the animal by biased head swing) and pirouette (bout of sharp turns generated by frequent reversals followed by omega turns) responses (see Nervous System-General Description) leading to impaired chemotaxis (Iino and Yoshida, 2009).
23
AIZR
Interneuron
Ventral ganglion in headRing Interneurons- SER-2; tyramine receptor (expresses two separate splice variants)
(Tsalik et al., 2003)
Integration of information from amphid sensory neurons; AIZL/R is one of the four first layer amphid interneuron pairs (AIA, AIB, AIY, and AIZ) that receive and process synaptic output from the amphid sensory neurons towards a behavioral response. Along with the second layer RIA and RIB interneurons, AIZ also receives heavy input from the first layer AIA, AIB, AIY interneurons and itself sends inputs to RIA and RIB. Additionally, it synapses directly onto SMB head motor neurons .
- Functions in thermotaxis; laser ablation of the AIZ pair makes the animals thermophilic (Mori and Ohshima, 1995). The thermal information sensed by AFD, AWC and ASI is processed in downstream interneurons AIY, AIZ, RIA, and also likely in AIA and AIB (the first three are considered as thermotaxis core interneurons) (Ma and Shen, 2012; Sasakura and Mori, 2012; Ardiel and Rankin, 2011; Beverly et al, 2011). In the original two-drive model of thermotaxis, activation of AIY neuron-signaling drive warm-seeking (thermophilic) movement, activation of AIZ neuron-signaling drive cold-seeking (cryophilic) movement, while the counterbalancing between the AIY and AIZ signals possibly through the RIA interneuron leads to execution of motor output in response to thermal signals (Ohnishi et al, 2011; Kuhara et al, 2008). However, there is evidence that a thermophilic drive at temperatures below cultivation temperature (Tc) does not exist and regulation of thermotaxis involves a more complex circuit (Ma and Shen, 2012; Ryu and Samuel, 2002). Around Tc animals track isothermally on a gradient and above Tc they display negative thermotaxis by regulating their turning frequency and run duration. It has been suggested that thermosensory signals from AFD, AWC and ASI converge on the AIB interneuron in negative thermotaxis, whereas the AIY/AIZ/RIA network may function mainly in isothermal tracking (Ma and Shen, 2012).
During associative learning between temperature and feeding state, starvation, a conditioning factor, downregulates AIZ activity through calcineurin/TAX-6, a calcium-activated phosphatase, leading to avoidance from Tc (Kuhara and Mori, 2006).
- Functions in locomotion: Laser ablations of AIZ, like AIB and AWC, cause decrease in reversals and lead to large increases in the duration of forward movement (Tsalik and Hobert, 2003; Iino and Yoshida, 2009). Since ablations of AIY and AIZ cause opposite behavioral phenotpes, the synapses made by AIY onto AIZ may be inhibitory. Alternatively, signals fom AIY to AIZ could be processed in a downstream neuron such as RIA (de Bono and Maricq 2005).
- Functions in chemotaxis: Laser ablations of AIZ cause defects in the weathervane (slow turning of the animal by biased head swing) and pirouette (bout of sharp turns generated by frequent reversals followed by omega turns) responses (see Nervous System-General Description) leading to impaired chemotaxis (Iino and Yoshida, 2009).
24
ALA
Interneuron
Dorsal ganglion of headHas two processes that branch from the anterior portion of the cell body, project into the left or right side of the nerve ring and then migrate as far as the tail on the lateral cords, adjacent to excretory canals. Also sends a process along dorsal cord.- FLP-7; FMRFamide-like peptide
(Li and Kim, 2008)
None yet reported, although described to have gap junctions in adult animals (MoW) - ACR-13 (LEV-8)
- ACR-4 (DES-2)
- SRA-10
(Van Buskirk and Sternberg, 2010; Towers et al., 2005; Troemel et al., 1995)
- Involved in inducing normal lethargus quiescence (i.e., cessation of pharyngeal pumping and locomotion during the lethargus periods (an EGF/LET-23-induced sleep-like state prior to molts). ALA neuron inhibits locomotion by inhibiting AVE, which normally functions to promote locomotion. Synapse between ALA and AVE is contacted by the CEPsh glia and CEPsh cells inhibit synaptic transmission from ALA to AVE, promoting locomotion.
(Mulcahy and Ient 2010; Van Buskirk and Sternberg, 2007)
25
ALML
Sensory neuron
Lateral in mid-bodyAnterior lateral microtubule cells, touch receptor neuronsGlutamate. Expresses VGluT, EAT-4 (Lee et al., 1999).Along with AVM and PLM, sense touch to the body and provide input to the command (inter) neurons (PVC, AVB, AVD, AVA) via both synaptic connections and gap junctions (Chalfie et al., 1985). The touch cells form gap junctions with agonist interneurons and chemical synapses with the antagonist interneurons. Hence, the anterior touch cells ALML/R and AVM form gap junctions with the backward movement interneuron AVD, but they provide synaptic input to the forward interneurons (AVB and PVC) (Kaplan and Driscoll, 1997; Goodman, 2006). See body touch circuit here.
Receptor Expression: Express MEC-2 (stomatin-like), MEC-4 (degenerin), MEC-10 (degenerin) which comprise part of a mechanosensory transduction channel (Huang et al., 1995; Lai et al., 1996; Huang and Chalfie, 1994). Express MEC-9 which is suggested to provide an extracellular attachment point for the mechanosensory channels in touch cells (Du et al., 1996) and MEC-6 which is part of the degenerin/epithelial Na+ channel complex (Chelur et al., 2002). Express glutamate receptor subunit, GLR-8 (Brockie et al., 2001). Express alpha subunits of a nicotinic acetylcholine receptor, DEG-3 and DES-2 (Treinin and Chalfie, 1995, Treinin. et al., 1998). Express the dopamine receptor DOP-1 (Tsalik et al., 2003, Sanyal et al., 2004).
26
ALMR
Sensory neuron
Lateral in mid-bodyAnterior lateral microtubule cells, touch receptor neuronsGlutamate. Expresses VGluT, EAT-4 (Lee et al., 1999).Along with AVM and PLM, sense touch to the body and provide input to the command (inter) neurons (PVC, AVB, AVD, AVA) via both synaptic connections and gap junctions (Chalfie et al., 1985). The touch cells form gap junctions with agonist interneurons and chemical synapses with the antagonist interneurons. Hence, the anterior touch cells ALML/R and AVM form gap junctions with the backward movement interneuron AVD, but they provide synaptic input to the forward interneurons (AVB and PVC) (Kaplan and Driscoll, 1997; Goodman, 2006). See body touch circuit here.
Receptor Expression: Express MEC-2 (stomatin-like), MEC-4 (degenerin), MEC-10 (degenerin) which comprise part of a mechanosensory transduction channel (Huang et al., 1995; Lai et al., 1996; Huang and Chalfie, 1994). Express MEC-9 which is suggested to provide an extracellular attachment point for the mechanosensory channels in touch cells (Du et al., 1996) and MEC-6 which is part of the degenerin/epithelial Na+ channel complex (Chelur et al., 2002). Express glutamate receptor subunit, GLR-8 (Brockie et al., 2001). Express alpha subunits of a nicotinic acetylcholine receptor, DEG-3 and DES-2 (Treinin and Chalfie, 1995, Treinin. et al., 1998). Express the dopamine receptor DOP-1 (Tsalik et al., 2003, Sanyal et al., 2004).
27
ALNL
Sensory neuron
TailNeurons associated with ALM, send processes into tailspikeAcetylcholine. Rand and Nonet, 1997.Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Express the dopamine receptor DOP-1 (Tsalik et al., 2003).- Soluble guanylate cyclase (sGC, e.g. GCY-35 and GCY-36)-expressing oxygen-sensing neurons (URX, AQR, PQR, SDQ, BDU, ALN, and PLN) mediate the avoidance of high O2 levels (Zimmer et al, 2009). URX appears to be the most important member of this group, since its activity is uniquely important for aggregation (Coates and de Bono, 2002), while for aerotactic behavior in an O2 gradient, URX is redundant with other sGC-expressing neurons (Chang et al., 2006).
28
ALNR
Sensory neuron
TailNeurons associated with ALM, send processes into tailspikeAcetylcholine. Rand and Nonet, 1997.Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Express the dopamine receptor DOP-1 (Tsalik et al., 2003).- Soluble guanylate cyclase (sGC, e.g. GCY-35 and GCY-36)-expressing oxygen-sensing neurons (URX, AQR, PQR, SDQ, BDU, ALN, and PLN) mediate the avoidance of high O2 levels (Zimmer et al, 2009). URX appears to be the most important member of this group, since its activity is uniquely important for aggregation (Coates and de Bono, 2002), while for aerotactic behavior in an O2 gradient, URX is redundant with other sGC-expressing neurons (Chang et al., 2006).
29
AQR
Sensory neuron
Head; right hand side near the posterior bulb of the pharynx
Postembryonically born neuron with rudimentary cilium. AQR and its ciliated dendrite are directly exposed to the pseudocoelomic body fluid. AQR axonal process projects into ring.Expresses orphan receptor (chemosensory/odorant?) guanylyl cyclase, GCY-32 (Yu et al., 1997). Expresses neuropeptide Y receptor like protein, NPR-1 which mediates social feeding behavior (Coates and de Bono , 2002). Expresses specific soluble guanylate cyclase homologue, GCY-35 which binds molecular oxygen and mediates oxygen sensation (Gray et al., 2004).Along with PQR, URX and AUA, AQR regulate social feeding (or aggregation on a bacterial lawn) and bordering (the accumulation of animals on the thickest part of a bacterial lawn) behavior since suppressing the activity of AQR, PQR and URX neurons inhibits social feeding (Coates and de Bono , 2002). AQR, PQR and URXfunction as sensors of environmental oxygen which is a quantitative regulator of social feeding (Gray et al., 2004). Decreases in oxygen leads to a dose-dependent suppression of social feeding behavior and bordering. Oxygen acts in parallel to NPR-1 in regulating social feeding behavior.
30
AS1
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).

- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
31
AS2
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
32
AS3
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
33
AS4
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
34
AS5
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
35
AS6
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
36
AS7
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
37
AS8
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
38
AS9
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
39
AS10
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
40
AS11
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, born postembryonically. Innervate dorsal muscles, no ventral counterpart, similar to VAn but receive additional synaptic input from AVB. Note that unlike AS1-10, the commissure of AS11 originates from the posterior process. All AS commissures to DC run from the right side of the body. Hermaphrodites and males exhibit some differences in ASn connectivity.Acetylcholine. Rand and Nonet, 1997. NLP-21; neuropeptide-like peptide
(Loer, 2010; von Stetina et al, 2007; Rand and Nonet, 1997).
- INX-3
- UNC-7
(Altun et al., 2009; Starich et al., 2009)
- ACR-14; nonalpha subunit of a nicotinic acetylcholine receptor
- ACR-15; alpha subunit of a nicotinic acetylcholine receptor
- LGC-46; ligand-gated ion channel
- UNC-63; alpha subunit of a levamisole-sensitive nicotinic acetylcholine receptor
(Wormbase; Altun, 2011; von Stetina et al, 2007; Fox et al, 2005).
41
ASEL
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from
ventral ganglion, make diverse synaptic connections in ring neuropil.
Express neuropeptide-like proteins, NLP-3, NLP-7, NLP-14 (Nathoo et al., 2001). Express the FMRFamide-related neuropeptide (FaRP), FLP-21 (Rogers et al., 2003).ASEL expresses orphan receptor (chemosensory/odorant?) guanylyl cyclases GCY-6 and GCY-7 and ASER expresses GCY-5 (Yu et al., 1997). Express mammalian capsaicin receptor-like protein OSM-9 (Colbert et al., 1997). Express neuropeptide Y receptor like protein, NPR-1 (Coates and de Bono , 2002).- Involved in chemotaxis to cAMP, biotin, Cl-, Na+ and lysine; ASEL is primarily sensitive to Na+ , whereas ASER is primarily sensitive to Cl- and K+ (Pierce-Shimomura et al., 2001; Bargmann and Horvitz, 1991; ). Opposite intacellular Ca++ transients are suggested to be generated in ASEL and ASER in response to changes in NaCl concentrations (Suzuki et al., 2004).
- ASH, ADL, ASK and ASE sensory neurons are responsible for the detection of certain chemical repellents. ASH plays a major role in this avoidance, whereas ADL, ASK and ASE play minor roles that are only evident when ASH is missing (Hiliard et al., 2002; Sambongi et al., 1999; Bargmann et al, 1990.) Mediate avoidance behavior from Cd2+ and Cu2+ (Sambongi et al., 1999)
- Functions in adaptive food-leaving behavior (leaving a food patch as it is becoming depleted to start foraging); the ASE neurons promote food-leaving behavior via a cGMP pathway as food becomes limited; in well-fed animals ASE neurons mediate chemoattraction to water-soluble cues as noted above. However, food withdrawal or gradual depletion of food turns this attraction to salt into repulsion. This switch in preference involves signaling by the DAF-2 insulin receptor in ASER as well as INS-1-dependent feedback from AIA to ASER (Milward et al., 2011; Tomioka et al., 2006)
42
ASER
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from
ventral ganglion, make diverse synaptic connections in ring neuropil.
Express neuropeptide-like proteins, NLP-3, NLP-7, NLP-14 (Nathoo et al., 2001). Express the FMRFamide-related neuropeptide (FaRP), FLP-21 (Rogers et al., 2003).ASEL expresses orphan receptor (chemosensory/odorant?) guanylyl cyclases GCY-6 and GCY-7 and ASER expresses GCY-5 (Yu et al., 1997). Express mammalian capsaicin receptor-like protein OSM-9 (Colbert et al., 1997). Express neuropeptide Y receptor like protein, NPR-1 (Coates and de Bono , 2002).- Involved in chemotaxis to cAMP, biotin, Cl-, Na+ and lysine; ASEL is primarily sensitive to Na+ , whereas ASER is primarily sensitive to Cl- and K+ (Pierce-Shimomura et al., 2001; Bargmann and Horvitz, 1991; ). Opposite intacellular Ca++ transients are suggested to be generated in ASEL and ASER in response to changes in NaCl concentrations (Suzuki et al., 2004).
- ASH, ADL, ASK and ASE sensory neurons are responsible for the detection of certain chemical repellents. ASH plays a major role in this avoidance, whereas ADL, ASK and ASE play minor roles that are only evident when ASH is missing (Hiliard et al., 2002; Sambongi et al., 1999; Bargmann et al, 1990.) Mediate avoidance behavior from Cd2+ and Cu2+ (Sambongi et al., 1999)
- Functions in adaptive food-leaving behavior (leaving a food patch as it is becoming depleted to start foraging); the ASE neurons promote food-leaving behavior via a cGMP pathway as food becomes limited; in well-fed animals ASE neurons mediate chemoattraction to water-soluble cues as noted above. However, food withdrawal or gradual depletion of food turns this attraction to salt into repulsion. This switch in preference involves signaling by the DAF-2 insulin receptor in ASER as well as INS-1-dependent feedback from AIA to ASER (Milward et al., 2011; Tomioka et al., 2006)
43
ASGL
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil.Express mammalian capsaicin receptor-like protein OSM-9 (Colbert et al., 1997). Express neuropeptide Y receptor like protein, NPR-1 (Coates and de Bono , 2002).Involved in chemotaxis to lysine, contribute to a residual chemotactic response to cAMP, biotin, Cl-, and Na+ after ASE is killed (Bargmann and Horvitz, 1991). Controls entry into dauer stage (Bargmann and Horvitz, 1991).
44
ASGR
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil.Express mammalian capsaicin receptor-like protein OSM-9 (Colbert et al., 1997). Express neuropeptide Y receptor like protein, NPR-1 (Coates and de Bono , 2002).Involved in chemotaxis to lysine, contribute to a residual chemotactic response to cAMP, biotin, Cl-, and Na+ after ASE is killed (Bargmann and Horvitz, 1991). Controls entry into dauer stage (Bargmann and Horvitz, 1991).
45
ASHL
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Glutamate. (Lee et al., 1999). Express neuropeptide-like proteins, NLP-3 and NLP-15 (Nathoo et al., 2001). Express the FMRFamide-related neuropeptide (FaRP), FLP-21 (Rogers et al., 2003).- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)-cation selective channel. Express TRPV channel proteins OSM-9 (Colbert et al., 1997) and OCR-2 (Tobin et al., 2002; de Bono M. et al., 2002). Express G protein-coupled serpentine receptors, SRA-6, SRB-6 (Troemel et al., 1995). Express neuropeptide Y receptor like protein, NPR-1 which mediates social feeding behavior (Coates and de Bono , 2002). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).- ASH plays a major role in avoidance responses to nose touch, hyperosmolarity (note: a mutation in the glr-1 (glutamate receptor subunit) gene which acts in synaptic targets of the ASH neurons, eliminates the response to nose touch but not to osmotic repellents), volatile repellent chemicals (1-octanol), heavy metals (Cd++ and Cu++), detergents/SDS, protons, alkaloids such as quinine (ASH is the main sensory neuron responsible for quinine detection and ASK plays a minor role) (de Bono & Villu Maricq, 2005; Bargmann, 2006; Hilliard et al, 2004; Hiliard et al., 2002; Hart et al., 1999; Sambongi et al., 2000; Sambongi et al., 1999; Troemel et al., 1995; Bargmann et al, 1990; Culotti & Russell 1978.)
- ASH and ADL are proposed to mediate social feeding behavior in response to repulsive cues (ablation of ASH and ADL abolishes social feeding behavior transforming social animals to solitary feeders) (de Bono M. et al., 2002.)
46
ASHR
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Glutamate. (Lee et al., 1999). Express neuropeptide-like proteins, NLP-3 and NLP-15 (Nathoo et al., 2001). Express the FMRFamide-related neuropeptide (FaRP), FLP-21 (Rogers et al., 2003).- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)-cation selective channel. Express TRPV channel proteins OSM-9 (Colbert et al., 1997) and OCR-2 (Tobin et al., 2002; de Bono M. et al., 2002). Express G protein-coupled serpentine receptors, SRA-6, SRB-6 (Troemel et al., 1995). Express neuropeptide Y receptor like protein, NPR-1 which mediates social feeding behavior (Coates and de Bono , 2002). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).- ASH plays a major role in avoidance responses to nose touch, hyperosmolarity (note: a mutation in the glr-1 (glutamate receptor subunit) gene which acts in synaptic targets of the ASH neurons, eliminates the response to nose touch but not to osmotic repellents), volatile repellent chemicals (1-octanol), heavy metals (Cd++ and Cu++), detergents/SDS, protons, alkaloids such as quinine (ASH is the main sensory neuron responsible for quinine detection and ASK plays a minor role) (de Bono & Villu Maricq, 2005; Bargmann, 2006; Hilliard et al, 2004; Hiliard et al., 2002; Hart et al., 1999; Sambongi et al., 2000; Sambongi et al., 1999; Troemel et al., 1995; Bargmann et al, 1990; Culotti & Russell 1978.)
- ASH and ADL are proposed to mediate social feeding behavior in response to repulsive cues (ablation of ASH and ADL abolishes social feeding behavior transforming social animals to solitary feeders) (de Bono M. et al., 2002.)
47
ASIL
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Express neuropeptide-like proteins, NLP-1, NLP-5, NLP-6, NLP-7, NLP-9, NLP-14, NLP-18, NLP-24, NLP-27 (Nathoo et al., 2001).Express G protein-coupled serpentine receptors, SRD-1, STR-2, STR-3 (Peckol et al.. 2001). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000). Gustatory sensory neurons. The main synpatic output of ASI is on to AIA interneuron.
- Involved in chemotaxis to lysine, contribute to a residual chemotactic response to cAMP, biotin, Cl-, and Na+ after ASE is killed (Bargmann and Horvitz, 1991).
- Controls entry into dauer stage (Bargmann and Horvitz, 1991).
- Functions in locomotion: Suppresses omega turns and reversals enhancing dispersal (Gray et al., 2005). After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later as reversal and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005).
- Thermosensor: After cultivation at a uniform temperature (Tc) with sufficient food, animals preferentially migrate to their cultivation temperature (Tc) when placed on a thermal gradient, and move isothermally at this temperature (Hedgecock and Russell 1975). Animals sense and record their Tc by AFD (major thermosensory), AWC and ASI neurons (Beverly et al, 2011; Kuhara et al., 2008; Biron et al., 2008). This Tc memory is plastic and can be reset upon cultivation at a different temperature (Hedgecock and Russell 1975). Unlike AFD neurons, which respond to thermal stimuli above Tc with continuous, graded calcium signals in a deterministic and highly reproducible way, ASI neurons exhibit temperature-induced stochastic Ca++ transients in a defined operating range with decreased responses close to or far from Tc (Beverly et al, 2011).
- Repression of pheromone (sexual) attraction in hermaphrodites. daf-7 mutation or ablation of the ASI neuron pair, which is the sole source of DAF-7/TGF-β inC. elegans, reveals sexual attraction (male-specific behavior) in hermaphrodites (White and Jorgensen, 2012).
48
ASIR
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Express neuropeptide-like proteins, NLP-1, NLP-5, NLP-6, NLP-7, NLP-9, NLP-14, NLP-18, NLP-24, NLP-27 (Nathoo et al., 2001).Express G protein-coupled serpentine receptors, SRD-1, STR-2, STR-3 (Peckol et al.. 2001). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000). Gustatory sensory neurons. The main synpatic output of ASI is on to AIA interneuron.
- Involved in chemotaxis to lysine, contribute to a residual chemotactic response to cAMP, biotin, Cl-, and Na+ after ASE is killed (Bargmann and Horvitz, 1991).
- Controls entry into dauer stage (Bargmann and Horvitz, 1991).
- Functions in locomotion: Suppresses omega turns and reversals enhancing dispersal (Gray et al., 2005). After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later as reversal and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005).
- Thermosensor: After cultivation at a uniform temperature (Tc) with sufficient food, animals preferentially migrate to their cultivation temperature (Tc) when placed on a thermal gradient, and move isothermally at this temperature (Hedgecock and Russell 1975). Animals sense and record their Tc by AFD (major thermosensory), AWC and ASI neurons (Beverly et al, 2011; Kuhara et al., 2008; Biron et al., 2008). This Tc memory is plastic and can be reset upon cultivation at a different temperature (Hedgecock and Russell 1975). Unlike AFD neurons, which respond to thermal stimuli above Tc with continuous, graded calcium signals in a deterministic and highly reproducible way, ASI neurons exhibit temperature-induced stochastic Ca++ transients in a defined operating range with decreased responses close to or far from Tc (Beverly et al, 2011).
- Repression of pheromone (sexual) attraction in hermaphrodites. daf-7 mutation or ablation of the ASI neuron pair, which is the sole source of DAF-7/TGF-β inC. elegans, reveals sexual attraction (male-specific behavior) in hermaphrodites (White and Jorgensen, 2012).
49
ASJL
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Express neuropeptide-like protein NLP-3 (Nathoo et al., 2001).Express mammalian capsaicin receptor-like protein OSM-9 (Colbert et al., 1997). Express G protein-coupled serpentine receptors, SRE-1 (Troemel et al., 1995). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000).Controls exit from dauer stage (Bargmann and Horvitz, 1991).
50
ASJR
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Express neuropeptide-like protein NLP-3 (Nathoo et al., 2001).Express mammalian capsaicin receptor-like protein OSM-9 (Colbert et al., 1997). Express G protein-coupled serpentine receptors, SRE-1 (Troemel et al., 1995). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000).Controls exit from dauer stage (Bargmann and Horvitz, 1991).
51
ASKL
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Glutamate. Expresses VGluT, EAT-4 (Lee et al., 1999). Express neuropeptide-like proteins, NLP-8, NLP-10, NLP-14 (Nathoo et al., 2001).- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)-cation selective channel
(Colbert et al., 1997).
- Express G protein-coupled serpentine receptors, SRA-7, SRA-9, SRG-2, SRG-8 (Troemel et al., 1995).
Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000).
- Involved in chemotaxis to lysine (Bargmann and Horvitz, 1991).
- ASH, ADL, ASK and ASE sensory neurons are responsible for the detection of certain chemical repellents. ASH plays a major role in this avoidance, whereas ADL, ASK and ASE play minor roles that are only evident when ASH is missing (Hiliard et al., 2002; Sambongi et al., 1999; Bargmann et al, 1990.) Mediate avoidance behavior from protons, detergents, alkaloids such as quinine -ASH is the main sensory neuron responsible for quinine detection while ASK plays a minor role (Hilliard et al, 2004).
- After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later, as reversals and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005).
- One of the three core sensory neurons (AWA, AWC, ASK) that are required for sexual-attraction in males (White and Jorgensen, 2012).
52
ASKR
Sensory neuron
Lateral ganglia of headAmphid neurons, single ciliated endings, project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil. Also take up FITC.Glutamate. Expresses VGluT, EAT-4 (Lee et al., 1999). Express neuropeptide-like proteins, NLP-8, NLP-10, NLP-14 (Nathoo et al., 2001).- OSM-9; TRPV (transient receptor potential channel, vanilloid subfamily; mammalian capsaicin receptor-like channel)-cation selective channel
(Colbert et al., 1997).
- Express G protein-coupled serpentine receptors, SRA-7, SRA-9, SRG-2, SRG-8 (Troemel et al., 1995).
Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000).
- Involved in chemotaxis to lysine (Bargmann and Horvitz, 1991).
- ASH, ADL, ASK and ASE sensory neurons are responsible for the detection of certain chemical repellents. ASH plays a major role in this avoidance, whereas ADL, ASK and ASE play minor roles that are only evident when ASH is missing (Hiliard et al., 2002; Sambongi et al., 1999; Bargmann et al, 1990.) Mediate avoidance behavior from protons, detergents, alkaloids such as quinine -ASH is the main sensory neuron responsible for quinine detection while ASK plays a minor role (Hilliard et al, 2004).
- After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later, as reversals and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005).
- One of the three core sensory neurons (AWA, AWC, ASK) that are required for sexual-attraction in males (White and Jorgensen, 2012).
53
AUAL
Interneuron
Lateral ganglia of headNeuron, process runs with amphid processes but lacks ciliated endingGlutamate. Expresses VGluT, EAT-4 (Lee et al., 1999).Express glutamate receptor subunit, GLR-4 (Brockie et al., 2001). Express neuropeptide Y receptor like protein, NPR-1 which mediates social feeding behavior (Coates and de Bono , 2002). Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Possibly express the dopamine receptor DOP-1 (Sanyal et al., 2004).Regulate social feeding behavior along with AQR, PQR and URX neurons (Coates and de Bono, 2002).
54
AUAR
Interneuron
Lateral ganglia of headNeuron, process runs with amphid processes but lacks ciliated endingGlutamate. Expresses VGluT, EAT-4 (Lee et al., 1999).Express glutamate receptor subunit, GLR-4 (Brockie et al., 2001). Express neuropeptide Y receptor like protein, NPR-1 which mediates social feeding behavior (Coates and de Bono , 2002). Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Possibly express the dopamine receptor DOP-1 (Sanyal et al., 2004).Regulate social feeding behavior along with AQR, PQR and URX neurons (Coates and de Bono, 2002).
55
AVAL
Interneuron
Lateral ganglia of headVentral cord interneuronFMRFamide (FLP-1). Express FMRFamide-like peptide, FLP-1 precursor (Nelson et al., 1998). Express the FMRFamide-related neuropeptide (FaRP), FLP-18 (Rogers et al., 2003).Express glutamate receptor subunits, GLR-1, GLR-2, GLR-4, GLR-5, NMR-1 and NMR-2 (Maricq et al., 1995; Brockie et al., 2001). Suggested to express GABA-A/glycine receptor-like protein GGR-3 (Fujiwara et al., 1996). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).Command interneuron. Functions as driver cell for backward locomotion. Drives backward movement of the animal along with touch modulator AVD neuron, AVE and A-type motor neurons.
56
AVAR
Interneuron
Lateral ganglia of headVentral cord interneuronFMRFamide (FLP-1). Express FMRFamide-like peptide, FLP-1 precursor (Nelson et al., 1998). Express the FMRFamide-related neuropeptide (FaRP), FLP-18 (Rogers et al., 2003).Express glutamate receptor subunits, GLR-1, GLR-2, GLR-4, GLR-5, NMR-1 and NMR-2 (Maricq et al., 1995; Brockie et al., 2001). Suggested to express GABA-A/glycine receptor-like protein GGR-3 (Fujiwara et al., 1996). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).Command interneuron. Functions as driver cell for backward locomotion. Drives backward movement of the animal along with touch modulator AVD neuron, AVE and A-type motor neurons.
57
AVBL
Interneuron
Lateral ganglia of headVentral cord interneuronExpress glutamate receptor subunits, GLR-1 and GLR-5 (Maricq et al., 1995; Brockie et al., 2001). Express G protein-coupled serpentine receptor, SRA-11 (Troemel et al., 1995). Suggested to express GABA-A/glycine receptor-like protein GGR-3 (Fujiwara et al., 1996). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).Command interneuron. Functions as driver cell for forward locomotion. Drives forward movement of the animal along with the touch modulator, PVC, and B-type motor neurons.
58
AVBR
Interneuron
Lateral ganglia of headVentral cord interneuronExpress glutamate receptor subunits, GLR-1 and GLR-5 (Maricq et al., 1995; Brockie et al., 2001). Express G protein-coupled serpentine receptor, SRA-11 (Troemel et al., 1995). Suggested to express GABA-A/glycine receptor-like protein GGR-3 (Fujiwara et al., 1996). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).Command interneuron. Functions as driver cell for forward locomotion. Drives forward movement of the animal along with the touch modulator, PVC, and B-type motor neurons.
59
AVDL
Interneuron
Lateral ganglia of headVentral cord interneuronExpress glutamate receptor subunits, GLR-1, GLR-2, GLR-5, NMR-1 and NMR-2 (Maricq et al., 1995; Brockie et al., 2001). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).Command interneuron. Functions as touch modulator for backward locomotion induced by head-touch. Drives backward movement of the animal along with driver cell AVA neuron, AVE and A-type motor neurons
60
AVDR
Interneuron
Lateral ganglia of headVentral cord interneuronExpress glutamate receptor subunits, GLR-1, GLR-2, GLR-5, NMR-1 and NMR-2 (Maricq et al., 1995; Brockie et al., 2001). Express UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997).Command interneuron. Functions as touch modulator for backward locomotion induced by head-touch. Drives backward movement of the animal along with driver cell AVA neuron, AVE and A-type motor neurons
61
AVEL
Interneuron
Lateral ganglia of headVentral cord interneuron, like AVD but outputs restricted to anterior cordFLP-1 precursor (Nelson et al., 1998).Express glutamate receptor subunits, GLR-1, GLR-2, GLR-5, NMR-1 and NMR-2 (Maricq et al., 1995; Brockie et al., 2001).Command interneuron. Drive backward movement of the animal along with AVA, AVD and A-type motor neurons.
62
AVER
Interneuron
Lateral ganglia of headVentral cord interneuron, like AVD but outputs restricted to anterior cordFLP-1 precursor (Nelson et al., 1998).Express glutamate receptor subunits, GLR-1, GLR-2, GLR-5, NMR-1 and NMR-2 (Maricq et al., 1995; Brockie et al., 2001).Command interneuron. Drive backward movement of the animal along with AVA, AVD and A-type motor neurons.
63
AVFL
Interneuron
Retrovesicular ganglion of headInterneuron
64
AVFR
Interneuron
Retrovesicular ganglion of headInterneuron
65
AVG
Interneuron
Retrovesicular ganglion of headVentral cord interneuronExpress glutamate receptor subunits, GLR-1, GLR-2, NMR-1 and NMR-2 (Maricq et al., 1995; Brockie et al., 2001). Expresses alpha subunit of a nicotinic acetylcholine receptor, DEG-3 (Treinin and Chalfie, 1995).Anterior guidepost neuron. AVG pioneers the right tract of the ventral nerve cord. It expresses UNC-6 at 3-fold embryo stage to provide a continuous UNC-6 (netrin)-labeled pathway restricted to the right tract of the nerve cord. The unilateral UNC-6 cue from AVG guides various paired axons growing from the nerve ring and lumbar ganglia to make rightward decussations as they enter the ventral nerve cord. UNC-6 also promotes bundling of axons within the cord itself. If the parent of AVG is ablated in wild-type embryos, interneurons and ventral cord motor neurons form several small fascicles rather than one tight bundle of right side tract and occasionally the right side axons shift to the left side (Wadsworth et al., 1996; Wadsworth and Hedgecock, 1986; Durbin 1987; Antebi et al., 1997).
66
AVHL
Interneuron
Lateral ganglia of headNeuron, mainly postsynaptic in ventral cord and presynaptic in the ringExpress glutamate receptor subunit GLR-4 (Brockie et al., 2001). Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Suggested to express GABA-A/glycine receptor-like protein GGR-1 (Fujiwara et al., 1996).
67
AVHR
Interneuron
Lateral ganglia of headNeuron, mainly postsynaptic in ventral cord and presynaptic in the ringExpress glutamate receptor subunit GLR-4 (Brockie et al., 2001). Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Suggested to express GABA-A/glycine receptor-like protein GGR-1 (Fujiwara et al., 1996).
68
AVJL
Interneuron
Lateral ganglia of headNeuron, synapses like AVHL/RPossibly glutamate (the neuron may be AIN instead). possibly expresses VGluT, EAT-4 (Lee et al., 1999).Express glutamate receptor subunit GLR-1 (Maricq et al., 1995).
69
AVJR
Interneuron
Lateral ganglia of headNeuron, synapses like AVHL/RPossibly glutamate (the neuron may be AIN instead). possibly expresses VGluT, EAT-4 (Lee et al., 1999).Express glutamate receptor subunit GLR-1 (Maricq et al., 1995).
70
AVKL
Interneuron
Ventral ganglion of headRing and ventral cord interneuronExpress FMRFamide-like peptide, FLP-1 precursor (Nelson et al., 1998).Express glutamate receptor subunit GLR-5 (Brockie et al., 2001).
71
AVKR
Interneuron
Ventral ganglion of headRing and ventral cord interneuronExpress FMRFamide-like peptide, FLP-1 precursor (Nelson et al., 1998).Express glutamate receptor subunit GLR-5 (Brockie et al., 2001).
72
AVL
Interneuron/motor neuron
Ventral ganglion in headRing and ventral cord interneuron and an excitatory GABAergic motor neuron for rectal muscles. Few synapsesGABA (Eastman et al., 1999)Defecation. AVL and DVB are excitatory GABAergic motor neurons for the enteric muscles (Avery and Thomas, 1997; McIntire et al., 1993). Together with DVB, activates expulsion muscle contraction (E.p or EMC), also required for anterior body contraction (aBoc) step in defecation motor program (DMP).
73
AVM
Sensory neuron
Lateral. Right side of the anterior half of the bodyAnterior ventral microtubule cell, touch receptorGlutamate. Expresses VGluT, EAT-4 (Lee et al., 1999).
74
AWAL
Sensory neuron
Lateral ganglia of headAmphid wing cells, neurons having ciliated sheet-like sensory endings closely associated with amphid sheathExpress 7-TM diacetyl receptor ODR-10 (Sengupta et al., 1996; Zhang et al., 1997). Express TRPV (mammalian capsaicin receptor-like channel) proteins OSM-9 (Colbert et al., 1997) and OCR-1 and OCR-2 (Tobin et al., 2002).Chemotaxis to diacetyl, pyrazine, trimethylthiazole (Bargmann et al., 1993). One of the three core sensory neurons (AWA, AWC, ASK) that are required for sexual-attraction in males (White and Jorgensen, 2012).
75
AWAR
Sensory neuron
Lateral ganglia of headAmphid wing cells, neurons having ciliated sheet-like sensory endings closely associated with amphid sheathExpress 7-TM diacetyl receptor ODR-10 (Sengupta et al., 1996; Zhang et al., 1997). Express TRPV (mammalian capsaicin receptor-like channel) proteins OSM-9 (Colbert et al., 1997) and OCR-1 and OCR-2 (Tobin et al., 2002).Chemotaxis to diacetyl, pyrazine, trimethylthiazole (Bargmann et al., 1993). One of the three core sensory neurons (AWA, AWC, ASK) that are required for sexual-attraction in males (White and Jorgensen, 2012).
76
AWBL
Sensory neuron
Lateral ganglia of headAmphid wing cells, neurons having ciliated sheet-like sensory endings closely associated with amphid sheathExpress neuropeptide-like proteins NLP-3 and NLP-9 (Nathoo et al., 2001).Express G protein-coupled serpentine receptor STR-1 (Troemel et al., 1997). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000). Suggested to express AEX-2, a 7-transmembrane domain protein with homology to the G protein-coupled receptor family (Mahoney et al., 2008).Avoidance from 2-nonanone, 1-octanol (Troemel et al., 1997).
77
AWBR
Sensory neuron
Lateral ganglia of headAmphid wing cells, neurons having ciliated sheet-like sensory endings closely associated with amphid sheathExpress neuropeptide-like proteins NLP-3 and NLP-9 (Nathoo et al., 2001).Express G protein-coupled serpentine receptor STR-1 (Troemel et al., 1997). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000). Suggested to express AEX-2, a 7-transmembrane domain protein with homology to the G protein-coupled receptor family (Mahoney et al., 2008).Avoidance from 2-nonanone, 1-octanol (Troemel et al., 1997).
78
AWCL
Sensory neuron
Lateral ganglia of headAmphid wing cells, neurons having ciliated sheet-like sensory endings closely associated with amphid sheath- Glutamate (AWC-AIA synapse is an inhibitory, glutamatergic synapse whose function is acutely modulated by NLP-1/NPR-11 signaling)
- NLP-1; neuropeptide-like protein
(Chalasani et al., 2010; Glauser and Goodman, 2010; Nathoo et al., 2001)
Express mammalian capsaicin receptor-like channel protein OSM-9 (Colbert et al., 1997). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000). Express G protein-coupled serpentine receptor, STR-2, asymmetrically (randomly on the right or left side) (Troemel et al., 1999). - AWC olfactory neurons are critical for chemotaxis to volatile odorants, (e.g. chemotaxis to benzaldehyde, butanone, isoamylalcohol, 2,3 pentanedione and 2,4,5 trimethylthiazole)
- They function in induction of local search behavior and promoting turns. After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later as reversal and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005).The AWC neurons synapse onto several interneurons including AIB and AIY, which enhance and suppress turning, respectively. The AWC neurons activate the AIB interneurons through AMPA-type glutamate receptors and inhibit AIY interneurons through glutamate-gated chloride channels. AWC are odor-OFF neurons that are activated by odor removal and inhibited in the continued presence of odors. On prolonged exposure to odorant (over 30 min), adaptation to AWC-sensed odors occurs, abrogating the chemotactic response. NLP-1 is required for these AWC-dependent food-evoked behaviors and odor adaptation (Chalasani et al., 2010; Chalasani et al., 2007; Gray et al., 2005; L'Etoile and Bargmann, 2000; Bargmann et al., 1993).
- Thermosensor: After cultivation at a uniform temperature (Tc) with sufficient food, animals preferentially migrate to their cultivation temperature (Tc) when placed on a thermal gradient, and move isothermally at this temperature (Hedgecock and Russell 1975). Animals sense and record their Tc by AFD (major thermosensory) and AWC and ASI neurons (Beverly et al, 2011; Kuhara et al., 2008; Biron et al., 2008). This Tc memory is plastic and can be reset upon cultivation at a different temperature (Hedgecock and Russell 1975). Activity of both the AFD and the AWC neurons is essential for the execution of thermotactic behaviors with high fidelity and precision. Upon sensation of a higher temperature than Tc, Ca++ concetration in AWC increases via production of cGMP by ODR-1 and activation of cGMP-dependent TAX-4 cation channel (Kuhara et al., 2008). Unlike AFD neurons, which respond to thermal stimuli above Tc with continuous, graded calcium signals in a deterministic and highly reproducible way, AWC neurons exhibit stochastic temperature-evoked Ca++ changes that are stimulus-correlated with different temporal kinetics at temperatures above and below Tc (Biron et al., 2008). AWC appears to get more active as temperatures rise or fall further from Tc. The AFD, AWC and ASI neurons seem to act in concert to increase turning rate when animals encounter higher temperatures than Tc on a gradient to move down the gradient toward colder temperatures near Tc (negative thermotaxis) (Biron et al., 2008). To track isotherms, animals do not actively pursue isothermal alignment, but once serendipitously aligned along an isotherm (at T=Tc), they track by suppressing turns (Luo et al., 2006). AWC neurons are relatively quiet around temperatures close to the Tc.
- One of the three core sensory neurons (AWA, AWC, ASK) that are required for sexual-attraction in males (White and Jorgensen, 2012).
79
AWCR
Sensory neuron
Lateral ganglia of headAmphid wing cells, neurons having ciliated sheet-like sensory endings closely associated with amphid sheath- Glutamate (AWC-AIA synapse is an inhibitory, glutamatergic synapse whose function is acutely modulated by NLP-1/NPR-11 signaling)
- NLP-1; neuropeptide-like protein
(Chalasani et al., 2010; Glauser and Goodman, 2010; Nathoo et al., 2001)
Express mammalian capsaicin receptor-like channel protein OSM-9 (Colbert et al., 1997). Express transmembrane guanylyl cyclase DAF-11 (Birnby et al., 2000). Express G protein-coupled serpentine receptor, STR-2, asymmetrically (randomly on the right or left side) (Troemel et al., 1999). - AWC olfactory neurons are critical for chemotaxis to volatile odorants, (e.g. chemotaxis to benzaldehyde, butanone, isoamylalcohol, 2,3 pentanedione and 2,4,5 trimethylthiazole)
- They function in induction of local search behavior and promoting turns. After animals are removed from bacterial food, they initiate a local search behavior consisting of reversals and deep omega-shaped turns. This is followed by dispersal ~30 min later as reversal and turns are suppressed. Local search behavior is triggered by AWC olfactory neurons, ASK gustatory neurons, and AIB interneurons while dispersal is promoted by ASI gustatory neurons and AIY interneurons (Gray et al., 2005).The AWC neurons synapse onto several interneurons including AIB and AIY, which enhance and suppress turning, respectively. The AWC neurons activate the AIB interneurons through AMPA-type glutamate receptors and inhibit AIY interneurons through glutamate-gated chloride channels. AWC are odor-OFF neurons that are activated by odor removal and inhibited in the continued presence of odors. On prolonged exposure to odorant (over 30 min), adaptation to AWC-sensed odors occurs, abrogating the chemotactic response. NLP-1 is required for these AWC-dependent food-evoked behaviors and odor adaptation (Chalasani et al., 2010; Chalasani et al., 2007; Gray et al., 2005; L'Etoile and Bargmann, 2000; Bargmann et al., 1993).
- Thermosensor: After cultivation at a uniform temperature (Tc) with sufficient food, animals preferentially migrate to their cultivation temperature (Tc) when placed on a thermal gradient, and move isothermally at this temperature (Hedgecock and Russell 1975). Animals sense and record their Tc by AFD (major thermosensory) and AWC and ASI neurons (Beverly et al, 2011; Kuhara et al., 2008; Biron et al., 2008). This Tc memory is plastic and can be reset upon cultivation at a different temperature (Hedgecock and Russell 1975). Activity of both the AFD and the AWC neurons is essential for the execution of thermotactic behaviors with high fidelity and precision. Upon sensation of a higher temperature than Tc, Ca++ concetration in AWC increases via production of cGMP by ODR-1 and activation of cGMP-dependent TAX-4 cation channel (Kuhara et al., 2008). Unlike AFD neurons, which respond to thermal stimuli above Tc with continuous, graded calcium signals in a deterministic and highly reproducible way, AWC neurons exhibit stochastic temperature-evoked Ca++ changes that are stimulus-correlated with different temporal kinetics at temperatures above and below Tc (Biron et al., 2008). AWC appears to get more active as temperatures rise or fall further from Tc. The AFD, AWC and ASI neurons seem to act in concert to increase turning rate when animals encounter higher temperatures than Tc on a gradient to move down the gradient toward colder temperatures near Tc (negative thermotaxis) (Biron et al., 2008). To track isotherms, animals do not actively pursue isothermal alignment, but once serendipitously aligned along an isotherm (at T=Tc), they track by suppressing turns (Luo et al., 2006). AWC neurons are relatively quiet around temperatures close to the Tc.
- One of the three core sensory neurons (AWA, AWC, ASK) that are required for sexual-attraction in males (White and Jorgensen, 2012).
80
BAGL
Sensory neuron
Anterior to the nerve ring in headNeuron, ciliated ending in head, no supporting cells, associated with ILsoExpress neuropeptide-like protein NLP-3 (Nathoo et al., 2001).Expresses orphan receptor (chemosensory/odorant?) guanylyl cyclase, GCY-33 (Yu et al., 1997).A neural circuit that includes BAG neurons mediate acute CO2 avoidance (Hallem and Sternberg, 2008; Bretscher et al., 2008)
81
BAGR
Sensory neuron
Anterior to the nerve ring in headNeuron, ciliated ending in head, no supporting cells, associated with ILsoExpress neuropeptide-like protein NLP-3 (Nathoo et al., 2001).Expresses orphan receptor (chemosensory/odorant?) guanylyl cyclase, GCY-33 (Yu et al., 1997).A neural circuit that includes BAG neurons mediate acute CO2 avoidance (Hallem and Sternberg, 2008; Bretscher et al., 2008)
82
BDUL
Interneuron
Lateral. Left and right sides of the anterior half of the bodyNeuron, process runs along excretory canal and into ring, unique darkly staining synaptic vesiclesExpress neuropeptide-like proteins, NLP-1 and NLP-15 (Nathoo et al., 2001).Express glutamate receptor subunit, GLR-8 (Brockie et al., 2001). Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003).Express neuropeptide-like proteins, NLP-1 and NLP-15 (Nathoo et al., 2001).
Function: Guidance for AVM process outgrowth during L1 stage (Walthall and Chalfie, 1988).
83
BDUR
Interneuron
Lateral. Left and right sides of the anterior half of the bodyNeuron, process runs along excretory canal and into ring, unique darkly staining synaptic vesiclesExpress neuropeptide-like proteins, NLP-1 and NLP-15 (Nathoo et al., 2001).Express glutamate receptor subunit, GLR-8 (Brockie et al., 2001). Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003).Express neuropeptide-like proteins, NLP-1 and NLP-15 (Nathoo et al., 2001).
Function: Guidance for AVM process outgrowth during L1 stage (Walthall and Chalfie, 1988).
84
CANL
Interneuron
Midbody, lateral sidesProcess runs along excretory canal, no synapses, essential for survivalMonoamine. Express VMaT, CAT-1 (Duerr et al., 1999). Express neuropeptide-like proteins, NLP-10 and NLP-15 (Nathoo et al., 2001).Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Suggested to express GABA-A/glycine receptor-like protein GGR-2 (Fujiwara et al., 1996).
85
CANR
Interneuron
Midbody, lateral sidesProcess runs along excretory canal, no synapses, essential for survivalMonoamine. Express VMaT, CAT-1 (Duerr et al., 1999). Express neuropeptide-like proteins, NLP-10 and NLP-15 (Nathoo et al., 2001).Express a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003). Suggested to express GABA-A/glycine receptor-like protein GGR-2 (Fujiwara et al., 1996).
86
CEPDL
Sensory neuron
HeadCephalic neurons, contain dopamineDopamine. Express TH, CAT-2 (Lints and Emmons, 1999).Express the dopamine receptor DOP-2 (Suo et al., 2003).Mechanosensory function in the head. When well-fed C. elegans hermaphrodites are washed clean of bacteria and then reintroduced to the bacterial lawn, they move more slowly than when transferred to an environment without bacteria. This behavior is described as "basal slowing response". The three classes of dopaminergic neurons (CEPs, ADEs, and PDEs) function redundantly to sense the mechanosensory stimulus from bacteria and mediate the motor circuit to control this behavioral change (Sawin et al., 2000).
87
CEPDR
Sensory neuron
HeadCephalic neurons, contain dopamineDopamine. Express TH, CAT-2 (Lints and Emmons, 1999).Express the dopamine receptor DOP-2 (Suo et al., 2003).Mechanosensory function in the head. When well-fed C. elegans hermaphrodites are washed clean of bacteria and then reintroduced to the bacterial lawn, they move more slowly than when transferred to an environment without bacteria. This behavior is described as "basal slowing response". The three classes of dopaminergic neurons (CEPs, ADEs, and PDEs) function redundantly to sense the mechanosensory stimulus from bacteria and mediate the motor circuit to control this behavioral change (Sawin et al., 2000).
88
CEPVL
Sensory neuron
HeadCephalic neurons, contain dopamineDopamine. Express TH, CAT-2 (Lints and Emmons, 1999).Express the dopamine receptor DOP-2 (Suo et al., 2003).Mechanosensory function in the head. When well-fed C. elegans hermaphrodites are washed clean of bacteria and then reintroduced to the bacterial lawn, they move more slowly than when transferred to an environment without bacteria. This behavior is described as "basal slowing response". The three classes of dopaminergic neurons (CEPs, ADEs, and PDEs) function redundantly to sense the mechanosensory stimulus from bacteria and mediate the motor circuit to control this behavioral change (Sawin et al., 2000).
89
CEPVR
Sensory neuron
HeadCephalic neurons, contain dopamineDopamine. Express TH, CAT-2 (Lints and Emmons, 1999).Express the dopamine receptor DOP-2 (Suo et al., 2003).Mechanosensory function in the head. When well-fed C. elegans hermaphrodites are washed clean of bacteria and then reintroduced to the bacterial lawn, they move more slowly than when transferred to an environment without bacteria. This behavior is described as "basal slowing response". The three classes of dopaminergic neurons (CEPs, ADEs, and PDEs) function redundantly to sense the mechanosensory stimulus from bacteria and mediate the motor circuit to control this behavioral change (Sawin et al., 2000).
90
DA1
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
91
DA2
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
92
DA3
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
93
DA4
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
94
DA5
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
95
DA6
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
96
DA7
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
97
DA8
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
98
DA9
Motor neuron
Body (ventral nerve cord)Ventral cord motor neurons, innervate dorsal musclesAcetylcholineExpresses UNC-8, a DEG/ENaC family member homologous to subunits of a candidate mechanically gated ion channel (Tavernarakis et al., 1997). DA9 expresses a splice variant of the tyramine receptor SER-2 (Tsalik et al., 2003; Goodman, 2006).Backward locomotion. DA's receive input from the driver interneuron AVA, and modulator interneurons AVD and AVE. They send output to VD neurons.
99
DB1
Motor neuron
Body (ventral nerve cord)|Ventral cord motor neurons, innervate dorsal muscles, reciprocal inhibitorAcetylcholineForward locomotion. DB's receive input from the command interneurons, AVB and PVC. They send output to VD neurons.
100
DB2
Motor neuron
Body (ventral nerve cord)|Ventral cord motor neurons, innervate dorsal muscles, reciprocal inhibitorAcetylcholineForward locomotion. DB's receive input from the command interneurons, AVB and PVC. They send output to VD neurons.
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Neurons
Neurotransmitters-Peptide Recep
ACH NeuroTransmitter
Serotonin Neurotransmitter
Dopamine Neurotransmitter
Tyramine Neurotransmitter
Octopamine Neurotransmitters
Glutamate Neurotransmitter
Gamma Neurotransmitter
Insulin-like peptides
FMRFamide-related peptides
Non-Insulin, non-FLP Peptides
flp neuropeptide
Sheet12