Descending

• Ijspeert, A. J. (2008). Central pattern generators for locomotion control in animals and robots: a review. Neural networks, 21(4), 642-653.

  • • ２．Neurobiology of CPGs:

      • • Central pattern generators (CPGs) are neural networks capable of producing coordinated patterns of rhythmic activity without any rhythmic inputs from sensory feedback or from higher controlcenters

        • リズム生成に感覚情報は不要．

        • This is in agreement with Grillner’s propositionthat CPGs are organized as coupled unit-burst elements with atleast one unit per articulation (i.e. per degree of freedom) inthe body (Grillner,1985).

        • Interestingly, simple signals are usually sufficient to induce activity in CPGs, as shown by the fictive locomotion experiments mentioned above. In many vertebrate animals, electrical stimula-tion of a specific region in the brain stem called Mesencephalic Lo-comotor Region (MLR) will induce locomotor behavior (Grillner,Georgopoulos,&Jordan,1997). The MLR is an important locomo-tor region that has descending pathways to the spinal cord via thereticular formations. Typically low-level stimulation2leads to slow(low frequency) movements, and high-level stimulation to faster(higher frequency) movements. The level of stimulation can there-fore modulate the speed of locomotion. Interestingly, MLR stimu-lation also induces automatic gait transition: in a decerebrated cat,increasing the stimulation leads to switches from walk to trot togallop (Shik,Severin,&Orlovsky,1966); in a decerebrated sala-mander increasing the stimulation leads to a switch from walkto swimming (Cabelguen,Bourcier-Lucas,&Dubuc,2003). Simi-lar gait transitions have been reported in other vertebrates (Grill-neret al.,1997). This demonstrates that CPGs are sophisticated circuits that can generate complex locomotor behaviors and even switch between very different gaits while receiving only simple in-put signals.3From a control point of view, CPGs therefore imple-ment some kind of internal model that ‘‘knows’’ which commandsignals need to be rhythmically produced to obtain a given speedof locomotion.

        • In the lamprey, the direction of locomotion can, similarly tovelocity, be modulated by simple variations of the stimulation applied to the MLR.

        • To summarize, the (vertebrate) locomotor system is organized such that the spinal CPGs are responsible for producing the basic rhythmic patterns, and that higher-level centers (the motor cortex,cerebellum, and basal ganglia) are responsible for modulating these patterns according to environmental conditions. Such a distributed organization presents several interesting features: (i) It reduces time delays in the motor control loop (rhythms are coordinated with mechanical movements using short feedback loops through the spinal cord). (ii) It dramatically reduces the dimensionality of the descending control signals. Indeed the control signals in general do not need to specify muscle activity but only modulate CPG activity. (iii) It therefore significantly reduces the necessary bandwidth between the higher-level centers and the spinal cord. CPGはリズムを生成する機能をもち，高次中枢は，環境に応じてこのリズムを調整する．利点：①運動制御の遅れを減らす，②下行性信号の次元を劇的に減らす（下行性信号は筋肉を直接コントロールする必要はない），③高次中枢と運動系の通信のBandwidthを劇的に減らす．

    • A properly implemented CPG model therefore reduces the dimensionality of the control problem such that higher-level controllers (or learning algorithms) do not need to directly produce multidimensional motor commands but only higher-level control signals. As discussed in Section 2, this is one of the most interesting features of biological CPGs. Related to this, CPG models typically produce smooth modulations of the produced trajectories even when the control parameters are abruptly changed (because the differential equations typically act as first or second order filters). See an example in Fig. 5. This property is useful for doing online trajectory generation that avoids possible damage in motors and gearboxes due to abrupt changes of motor commands.7

    • Design methodologies for CPGs (Chap 5)

      • • 4. Theeffect of input signals, i.e. how control parameters can modulateimportant quantities such as the frequency, amplitude, phaselags (for gait transition), or waveforms (e.g. for independentlyadjusting swing and stance phases)

    • Many researchers now propose that animal motor control is based on the combination of motor primitives, i.e. complex movements are generated by combining a finite set of simpler elementary movements (Flash and Hochner, 2005, Thoroughman and Shadmehr, 2000, Todorov, 2004, Tresch et al., 2002). Motor primitives (and related concepts such as muscle synergies, force fields, and motor schemas) are seen like elementary controllers that produce specific movements under the control of a few open control parameters. Experiments on decerebrated and spinalized animals indicate that, like CPGs, many of these motor primitives are implemented at a low level in the vertebrate central nervous system, namely in the brainstem and the spinal cord (Bizzi et al., 2000, Grillner, 2006, Stein and Smith, 1997, Tresch et al., 2002, Whelan, 1996). The interesting features of CPGs discussed in Section 2, e.g. in terms of the dimensionality of the control signals, can indeed also be found in discrete pattern generators, see the force field concept identified by Bizzi et al. (2000). CPGs should therefore be seen as particular movement primitives that can be activated together with others. From a robotics point of view, the idea of using motor primitives for constructing controllers for complex motor skills is appealing and is attracting a growing number of researchers (Ijspeert et al., 2003, Mussa-Ivaldi, 1997, Schaal and Schweighofer, 2005, Todorov et al., 2005).

• Loeb, G. E. (2001). Learning from the spinal cord. The Journal of physiology, 533(1), 111-117. 自由度を減らす，操り人形の例．

• Giszter, S. F. (2015). Motor primitives—new data and future questions. Current opinion in neurobiology, 33, 156-165.

• Gallego, J. A., Perich, M. G., Miller, L. E., & Solla, S. A. (2017). Neural manifolds for the control of movement. Neuron, 94(5), 978-984.

• Flash, T., & Bizzi, E. (2016). Cortical circuits and modules in movement generation: experiments and theories. Current opinion in neurobiology, 41, 174-178.

Borgmann A, Buschges A (2015) Insect motor control: methodological advances, descending control and inter-leg coordination on the move. Curr Opin Neurobiol. 33:8-15. Production of behavior relies on individual descending command like interneurons, but only for robuts, innate, and/or species specific behavior (except back up). How about transitional aspects e.g. direction/speed.

Tschida K, Bhandawat V (2015) Activity in descending dopaminergic neurons represents but is not required for leg movements in the fruit fly Drosophila. Physiol Rep 3:e12322.

Grillner S, El Manira A (2015) The intrinsic operation of the networks that makes us locomote. Curr Opin Neurobiol 31:244-249.

• *Steeves JD, Sholomenko GN, Webster DM (1987) Stimulation of the pontomedullary reticular formation initiates locomotion in decerebrate birds. Brain Res 401:205-212. MLR stimulation drives flight-like muscle activity.

• Shik M.L., Severin F.V., Orlovsky G.N. (1966) Control of walking and running by means of electrical stimulation of the midbrain. Biophysics 11: 756-765.

• Progressin Neurobiology V 49, 481 515,1996 C ~ 1 ElsevierS Ltd.Allrightsr P G B 0 PII: S0301-0082(%)00028-7 CONTROL OF LOCOMOTION IN THE DECEREBRATE CAT PATRICK J. WHELAN*

• Brain Research Volume 300, Issue 2, 23 May 1984, Pages 357–361 Cover image Activation of ‘fictive swimming’ by electrical microstimulation of brainstem locomotor regions in an in vitro preparation of the lamprey central nervous system Andrew D. McClellan, Sten Grillner

• DN019 Fig.6C Oh, Y., Yoon, S. E., Zhang, Q., Chae, H. S., Daubnerová, I., Shafer, O. T., ... & Kim, Y. J. (2014). A homeostatic sleep-stabilizing pathway in Drosophila composed of the sex peptide receptor and its ligand, the myoinhibitory peptide. PLoS Biol,12(10), e1001974.

http://www.humanneurophysiology.com/desendingmotorpathways.htm

Flycircuit DB

• fru-M-000012

Descending neurons gross anatomy

• Lemon, R. N. (2008). Descending pathways in motor control. Annu. Rev. Neurosci., 31, 195-218. Ventromedial brainstem pathway: a bilateral postural control system for head, neck, trunk, and proximal limb movements / Dorsolateral brainstem pathway / to provide additional capacity for flexion-biased movements involving more distal limb segments, the elbow and wrist.

• Coggshall JC, Boschek CB Buchner SM (1973) Preliminary investigations on a pair of giant fibers in the central nervous system of dipteran flies. Z Naturforsch 28c:783-784. neck connectiveには数千が走行．

• Strausfeld NJ, Bassemir U, Singh RN, Bacon JP (1984) Organizational principles of outputs from dipteran brains. J Insect Physiol 30:73-93. Basic organization of DNs. Concept for ‘parallel projecting descending neuron’ (PDN). Description about direct and indirect pathway. 電顕データに基づき，DNの軸索間の相互シナプスについて記述．MB output onto the Descending neurons (Fig. 15) -NOT describe synaptic connection by electron microcope, but schematic (Fig.16)

• Strausfeld NJ, Lee JK (1991) Neuronal basis for parallel visual processing in the fly. Visual Neurosci 7:13-33. Texas red fills demonstrate 300 pairs of DNs originating in the brain of which 120 arise from the deutocerebrum (n prep). 視覚系に2つの主要な経路．Lobula-Optic glomeruli-thoracic ganglia 背側（主に飛翔神経叢に対応），Lobula plate-Posterior slope-thoracic ganglia 腹側（主に歩脚神経叢に対応）．霊長類magnocellular and parvocellular pathwayと比較．

• Burdohan JA, Comer CM (1996) Cellular organization of an antennal mechanosensory pathway in the cockroach, Periplaneta americana. J Neurosci 16:5830-5843. Meso, metaの間からバックフィル，細胞体は40-45．

• Staudacher E (1998) Distribution and morphology of descending brain neurons in the cricket Gryllus bimaculatus. Cell Tissue Res 294:187-202. Backfill，脳（食道上神経節）の細胞体は約200個．

• Heinrich R (2002) Impact of descending brain neurons on the control of stridulation, walking, and flight in orthoptera. Microsc Res Tech 56:292-301.

• Okada R, Sakura M, Mizunami M (2003) Distribution of dendrites of descending neurons and its implications for the basic organization of the cockroach brain. J Comp Neurol 458:158-174. Backfill，脳（食道上神経節）の細胞体は235-284個．

• Cardona A, Larsen C, Hartenstein V (2009) Neuronal fiber tracts connecting the brain and ventral nerve cord of the early Drosophila larva. J Comp Neurol 515:427-440. ショウジョウバエ幼虫のVNCにDye injection，脳での細胞体・樹状突起の分布を観察．

• Børø S (2012) Morphological characterization of descending interneurons and determination of output areas in the brain of the moth Heliothis virescens. PhD Thesis, Norwegian University of Science and Technology. Visually comparing the neural arborization of DNs in the female and male, revealed that the male LAL are more heavily innervated by DN dendrites than the female / The staining of the LALs in the female brain was relatively weak, suggesting the LAL to be less innervated by DNs in females. In addition, was the awareness that LAL is involved in sex-specific pheromone information processing and behavior as shown in moth species, including H.virescens.

Descending neurons innervating antennal lobe

• Stocker RF, Lienhard MC, Borst A, Fischbach KF (1990) Neuronal architecture of the antennal lobe in Drosophila melanogaster. Cell Tissue Res 262:9-34. “A thoracic relay interneuron (TI) with extensive arborizations in the ventral half of the antennal lobe and the posterior-lateral brain, as well as a process leading into the thoracic ganglion”. DN006

• Nässel DR, Cantera R, Karlsson A (1992) Neurons in the cockroach nervous system reacting with antisera to the neuropeptide leucokinin I. J Comp Neurol 322:45-67. ALにdendriteを持つDN．

• Tanaka NK, Endo K, Ito K (2012) Organization of antennal lobe-associated neurons in adult Drosophila melanogaster brain. J Comp Neurol 520:4067-4130. AL-DN1 innervates AVLP, ASLP, PSLP, ASLP, RN, SCL, and SEG. DN004

Descending neurons from optic lobe

• Nässel DR, Strausfeld NJ (1982) A pair of descending neurons with dendrites in the optic lobes projecting directory to thoracic ganglia of dipterous insects. Cell Tissue Res 226:355-362. Lobula descending neuron (LDN) mainly described in Musca domestica. Also described in Calliphora erythocephala & Drosophila melanogaster. DN058

DCMD

• O'Shea M, Rowell CHF, Williams JLD (1974) The anatomy of a locust visual interneurone; The descending contralateral movement detector. J Exp Biol 60:1-12. Morphology of DCMD.DCMD run through dorsal-medial part of the VNC. DN032 has branches in the SEG, but DN032b doesn't (same to DCMD in locust). DN032b extended fiber to T2, T3 neuromere, which is similar to locust DCMD (3 main branches in T2/T3, only single branch in T1). The second branch projects ventrally and towards the midline, and then sends to two fine axons to the ipsi, and contralateral margins of the ganglion. These terminate laterally, just anterior to the exit of the main leg nerve, in a small zone of arborization. "neural variation is relatively great, in so far as it can be intuitively judged; that is, the structure of the DCMD, were it immediately apparent on inspection of the animal, would not be a good taxonomic character compared with many other morphological feature." "variation in individual neurons is the morphological correlate of the behavioral variance of the population, that such a range of behavioural capacity is a significant evolutionary asset, which selection could act to preserve and increase."

Descending neurons posterior slope

• Guy RG, Goodman LJ, Mobbs PG (1979) Visual interneurons in the bee brain: Synaptic organization and transmission by graded potentials. J Comp Physiol A 134:253-264.

• Strausfeld NJ, Bassemir UK (1985) Lobula plate and ocellar interneurons converge onto a cluster of descending neurons leading to neck and leg motor neuropil in Calliphora erythrocephala. Cell Tissue Res 240:617-640. DN013, DN029

• Milde JJ, Strausfeld NJ (1986) Visuo-motor pathways in arthropods: Giant motion-sensitive neurons connect compound eyes directory to neck muscles in blowflies (Calliphora erythrocephala). Naturwissenschaften 73:151-154.

• Haag J, Borst A (2005) Dye-coupling visualizes networks of large-field motion-sensitive neurons in the fly. J Comp Physiol A 191:445-454.

• Haag J, Wertz A, Borst A (2007) Integration of lobula plate output signals by DNOVS1, an identified premotor descending neuron. J Neurosci 27:1991-2000. DNOVS1(Descending neurons of the ocellar and vertical system) in Calliphora vicina.

• Wertz A, Borst A, Haag J (2008) Nonlinear integration of binocular optic flow by DNOVS2, a descending neuron of the fly. J Neurosci 28:3131-3140. DNOVS2 in Calliphora vicina.

• Wertz A, Haag J, Borst A (2009) Local and global motion preferences in descending neurons of the fly. J Comp Physiol A 195:1107-1120.

• Wertz A, Gaub B, Plett J, Haag J, Borst A (2009) Robust coding of ego-motion in descending neurons of the fly. J Neurosci 29:14993-15000.

• Hung YS, van Kleef JP, Stange G, Ibbotson MR (2013) Spectral inputs and ocellar contributions to a pitch-sensitive descending neuron in the honeybee. J Neurophysiol 109:1202-1213.

VLP and Other dorsal DNs

• Strausfeld NJ, Gronenberg W (1990) Descending neurons supplying the neck and flight motor of Diptera: Organization and neuroanatomical relationships with visual pathways. J Comp Neurol 302:954-972. DN形状をBilateral, Heterolateral, Homolateralに分類．形状タイプの構成とDendiritc Clusterが対応する．浅い方から，(1) Deeper layer ではBilateralがDominant (2) これより浅いDC2-4ではHeterolateral, Homolateralが寄与．(3) DC1, exclusively homolateral, including DNOVSs.

• Gronenberg W, Strausfeld NJ (1990) Descending neurons supplying the neck and flight motor of Diptera: Physiological and anatomical characteristics. J Comp Neurol 302:973-991.

• Milde JJ, Strausfeld (1990) Cluster organization and response characteristics of the giant fiber pathway of the blowfly Galliphora erythrocephala. J Comp Neurol 294:59-75. 8 DNs that contribute to a DN cluster located ventrally in the lateral deutocerebrum, an area interposed between the ventral antennal lobes and the laterally disposed optic lobes. Similarity with Drosophila DNs (i.e GDN/giant fiber, lateral giant/DN133, inferior giant/DN148, ipsilateral small/DN001, contralateral giant/DN032b). ロビュラからのColAニューロンが収束するOptic glomerulusに，樹状突起分枝が重複するDNのグループが存在する(Cluster organization，特にGDNのグループをGDNC)．最も顕著なGiant Descending Neuron（GDN）に加えておそらく電気シナプスで共役した5つのDN，2つの小型のDNが同じ領域に樹状突起を持ち，主に歩脚神経叢へ投射する．中心複合体からOptic glomeruliへの介在ニューロンなど．GDNCはロビュラ，ammc，Ascending neurons (Simialr to DN002) の入力を受ける．DN axons occupy the upper and lateral two thirds of the neck connective's cross section.

• Gronenberg W, Strausfeld NJ (1991) Descending pathways connecting the male-specific visual system of flies to the neck and flight motor. J Comp Physiol A 169:413-426.

• Gronenberg W, Strausfeld NJ (1992) Premotor descending neurons responding selectively to local visual stimuli in flies. J Comp Neurol 316:87-103.

• Gronenberg W, Milde JJ, Strausfeld NJ (1995) Oculomotor control in Calliphorid flies: organization of descending neurons to neck motor neurons responding to visual stimuli. J Comp Neurol 361:267-284. 2 small SMP DN

• Ruta V, Datta SR, Vasconcelos ML, Freeland J, Looger LL, Axel R (2010) A dimorphic pheromone circuit in Drosophila from sensory input to descending output. Nature 468:686-690. First recording from Drosophila DN， DN105?

• Träger U, Homberg U (2011) Polarization-sensitive descending neurons in the locust: Connecting the brain to thoracic ganglia. J Neurosci 31:2238-2247.

• von Philipsborn AC, Liu T, Yu JY, Masser C, Bidaye SS, Dickson BJ (2011) Neuronal control of Drosophila courtship song. Neuron 69:509-522.

• Mu L, Bacon JP, Ito K, Strausfeld NJ (2014) Responses of Drosophila giant descending neurons to visual and mechanical stimuli. J Exp Biol 217:2121-2129.

• Stubblefield, G. T., & Comer, C. M. (1989). Organization of giant interneuron projections in thoracic ganglia of the cockroach Periplaneta americana. Journal of Morphology, 200(2), 199-213.

Neck motor neurons

• Strausfeld NJ, Seyan HS (1985) Convergence of visual, haltere, and prosternai inputs at neck motor neurons of Calliphora erythrocephala. Cell Tissue Res 240:601-615. 400-500 fiber for haltere afferent

• Milde JJ, Seyan HS, Strausfeld NJ (1987) The neck motor system of the fly Calliphora erythrocephala. J Comp Physiol A 160:225-238.

Descending neurons from lateral accessory lobe

• Olberg RM (1986) Identified target-selective visual interneurons descending from the dragonfly brain. J Comp Physiol A 159:827-840. Visual control of flight orientation in dragonfly. (LAL DN is mentioned by Zorovic 2013)

• Kanzaki R, Shibuya T (1986) Descending protocerebral neurons related to the mating dance of the male silkworm moth. Brain Res 377:378-382. First description of long-lasting activity for LAL DNs.

• Kanzaki R, Arbas EA, Hildebrand JG (1991) Physiology and morphology of descending neurons in pheromone-processing olfactory pathways in the male moth Manduca sexta. J Comp Physiol A 169:1-14. DN innervating LAL showing long lasting activity in response to sex pheromone.

• Homberg U (1994) Flight-correlated activity changes in neurons of the lateral accessory lobes in brain of the locust Schistocereca gregaria. J Comp Physiol A 175:597-610. Related to flight control in locust. Responsive to air stimulation during fictive flight.

• Kanzaki R, Ikeda A, Shibuya T (1994) Morphological and physiological properties of pheromone-triggered flipflopping descending interneurons of the male silkworm moth, Bombyx mori. J Comp Physiol A 175:1-14. Extracellular recording from neck connective with suction electrode. Dye-fill indicate that the neurons with 'flip-flop' signal have dendritic innervation to the LAL.

• Mishima T, Kanzaki R (1999) Physiological and morphological characterization of olfactory descending interneurons of the male silkworm moth, Bombyx mori. J Comp Physiol A 184:143-160.

• Staudacher E (2001) Sensory responses of descending brain neurons in the walking cricket, Gryllus bimaculatus. J Comp Physiol A 187:1-17. DBNc5-5 show similar morphology to B-DC1. Activity seems to be correlated with forward velocity and rotational velocity.

• Wada S, Kanzaki R (2005) Neural control mechanisms of the pheromone-triggered programmed behavior in male silkmoths revealed by double-labeling of descending interneurons and a motor neuron. J Comp Neurol 484:168-182.

• Yamagata N, Nishino H, Mizunami M (2007) Neural pathways for the processing of alarm pheromone in the ant brain. J Comp Neurol 505:424-442. One type of pheromone-sensitive DN innervating LAL exhibit long-lasting activity.

• Yu JY, Kanai MI, Demir E, Jefferis GS, Dickson BJ (2010) Cellular organization of the neural circuit that drives Drosophila courtship behavior. Curr Biol 20:1602-1614. One intersection between GAL4/fruitless specify two types of LAL DNs (aSP3, aDT8). DN099

• Zorović M, Hedwig B (2011) Processing of species-specific auditory patterns in the cricket brain by ascending, local, and descending neurons during standing and walking. J Neurophysiol 105:2181-2194. Sound response of LAL DN (B-DC1(5)). DN095

• Zorović M, Hedwig B (2013) Descending brain neurons in the cricket Gryllus bimaculatus (de Geer): auditory responses and impact on walking. J Comp Physiol A 199:25-34. B-DI1(1) for ipsilateral innervation, B-DC1(5) for contralateral innervation. Activity of most DNs are correlated with forward velocity. B-DC1(5) activity is correlated with steering velocity. Activation of both B-DI1(1) and B-DC1(5) elicit walking. DN003/DN095

• Bidaye SS, Machacek C, Wu Y, Dickson BJ (2014) Neuronal control of Drosophila walking direction. Science 344:97-101. Two pairs of Moon-walker DNs have innervation into the LAL.

Possible homologous neurons to i5, c5-2

• Brodfuehrer PD, Hoy RR (1990) Ultrasound sensitive neurons in the cricket brain. J Comp Physiol A 166:651-662. DBIN7 show similar morphology to B-DC1, which show phasic/tonic activity.

• Williams JLD (1975) Anatomical studies of the insect central neervous system: a ground-plan of the midbrain and an introduction (Orthoptera). J Zool (Lond) 176:67-86. similar to i-5

LAL-PS connectivity

• Heinze S, Reppert SM (2011) Sun compass integration of skylight cues in migratory monarch butterflies. Neuron 69:345-358.

POTu

The most posterior (n-dorsal ) optic glomerulus of the brain; it has a distinct and prominent structure in e.g. locusts, crickets, cockroaches, and monarch butterfly (§ not prominent in Drosophila). (Ito et al., 2014). The posterior optic tubercle is the posteriormost optic glomerulus of the lateral brain, behind the PLP and ventral to the MB calyx. It has a prominent discrete structure in many locusts, crickets and cockroaches (Homberg, 1991; Homberg et al., 1991) as well as in monarch butterflies (Heinze and Reppert, 2012), but appears to be just one of several similar optic glomeruli of the PLP in Drosophila. In species with a prominent POTU, it can be identified by labeling large glomerulus-like arborizations emerging from the

posterior optic commissure as well as by tracing tangential neurons of the PB.

• Homberg U., Würden S. (1997) Movement-sensitive, polarization-sensitive, and light-sensitive neurons of the medulla and accessory medulla of the locust, Schihstocerca gregaria. J Comp Neurol 386:329346. input from the accessory medulla, TB1/TB2 connect PB and POTu.

• Helfrich-Förster C., Stengl M., Homberg U. (1998) Organization of the circadian system in insects. Chronobiol. Int 15:567-594. POTu receives internal circadian clock in cockroaches and flies

Target selective descending interneurons

• Olberg RM (1986) Identified target-selective visual interneurons descending from the dragonfly brain. J Comp Physiol A 159:827-840.

• Adelman TL, Bialek W, Olberg RW (2003) The information content of receptive fields. Neuron 40:823-833. Neuron 40:823-833.

• Frye MA, Olberg RM (1995) Visual receptive field properties of feature detecting neurons in the dragonfly. J Comp Physiol A 177:569-576.

• Gonzalez-Bellido PT, Peng H, Yang J, Georgopoulos AP, Olberg RM (2013) Eight pairs of descending visual neurons in the dragonfly give wing motor centers accurate population vector of prey direction. Proc Natl Acad Sci USA 110:696-701.

Mechanosensory descending neurons

• Burdohan JA, Comer CM (1996) Cellular organization of an antennal mechanosensory pathway in the cockroach, Periplaneta Americana. J Neurosci 16:5830-5843.

• Ache JM, Dürr V (2013) Encoding of near-range spatial information by descending interneurons in the stick insect antennal mechanosensory pathway. J Neurophysiol 110:2099-2112.

Aminergic/peptidergic descending neurons

• Nӓssel DR, Elekes K (1985) Serotonergic terminals in the neural sheath of the blowfly nervous system: Electron microscopical immunocytochemistry and 5,7-dihydroxytriptamine labeling. Neuroscience 15:293-307. 5HT DN in blowfly.

• Melcher C, Rankratz MJ (2005) Candidate gustatory interneurons modulating feeding behavior in the Drosophila brain. PLoS Biol 3:e305. 神経ペプチド hugin, Fig5b, one neuron in hugS3-GAL4 line resembles to DN138.

• Nӓssel DR, Homberg U (2006) Neuropeptide in interneurons of the insect brain. Cell Tissue Res 326:1-24. FMRF amide positive DN (SP1, DN115)

• Busch S, Selcho M, Ito K, Tanimoto H (2009) A map of octopaminergic neurons in Drosophila brain. J Comp Neurol 513:643-667. Octopaminergic cells in Drosophila. OA-VPM1 shows the same morphology to aDT8 in Yu et al. 2010 (DN099). DN047

• Alekseyenko OV, Lee C, Kravitz EA (2010) Targeted manipulation of serotonergic neurotransmission affects the escalation of aggression in adult male Drosophila melanogaster. PLoS One 5:e10806. TRH Gal4 line contains two big cells. At least one of them is descending neuron that has ‘heavy’ innervation into ventral nerve cord. DN052a/DN052b

• Certel SJ, Leung A, Lin CY, Perez P, Chiang AS, Kravitz EA (2010) Octopamine neuromodulatory effects on a social behavior decision-making network in Drosophila male. PLoS One 5:e13248. Intersection between FruM and tdc4 specify several neurons involved in social behavior, including OA-VPM1 (Busch et al. 2009). DN099

• de Haro M, Al-Ramahi I, Benito-Sipos J, López-Arias B, Dorado B, Veenstra JA, Herrero P (2010) Detailed analysis fo leucokinin-expressing neurons and their candidate functions in the Drosophila nervous system. Cell Tissue Res 339:321-336. Suboesophageal leucokinin-expressing neurons (SELK). GMR22C10

Ocellar DN (LD neurons) Direct innervation to ocellar ganglion DN118 in Drosophila

• Heinzeller T (1976) Second-order ocellar neurons in the brain of the honeybee (Apis mellifera). Cell Tissue Res 171:191-199.

• Pan KC, Goodman LJ (1977) Ocellar projections within the central nervous system of the worker honey bee, Apis mellifera. Cell Tissue Res 176:505-527.

• Goodman LJ (1981) Organization and physiology of the insect dorsal ocellar system. In Handbook of Sensory Physiology, VII/6C, (H. Autrum ed) Berlin, Heidelberg, New York, Springer, pp201-286.

• Milde JJ (1984) Ocellar interneurons in the honeybee - structure and signals of L-neurons. J Comp Physiol A 154:683-693.

• Milde JJ, Homberg U (1984) Ocellar interneurons in the honeybee: characteristics of spiking L-neurons. J Comp Physiol A 155:151-160.

• Hung YS, van Kleef JP, Stange G, Ibbotson MR (2013) Spectral inputs and ocellar contributions to a pitch-sensitive descending neuron in the honeybee. J Neurophysiol 109:1202-1213.

• Hung YS, Ibbotson MR (2014) Ocellar structure and neural innervation in the honeybee. Front Neuroanat 8:6. Honeybee 5 pairs of ocellar DNs.

• Ibbotson MR (2001) Evidence for velocity-tuned motion-sensitive descending neurons in the honeybee. Proc Roy Soc Lond B 268:2195-2201.

Visual computation in TSDNs

• O’Carroll DC, Bidwell NJ, Laughlin SB, Warrant EJ (1996) Insect motion detectors matched to visual ecology. Nature 382:63-66.

• Bolzon DM, Nordstrom K, O’Carroll DC (2009) Local and large-range inhibition in feature detection. J Neurosci 29:14143-14150.

• Barnett PD, Nordström K, O’Carroll DC (2010) Motion adaptation and the velocity coding of natural scenes. Curr Biol 20:994-999.

• Wiederman SD, O’Carroll DC (2011) Discrimination of features in natural scenes by a dragonfly neuron. J Neurosci 31:7141-7144.

• Wiederman SD, O’Carroll DC (2013) Selective attention in an insect visual neuron. Curr Biol 23:156-161.

Zebrafish

• Hagglund M, Borgius L, Dougherty K, Kiehn O (2010) Activation of groups of excitatory neurons in the mammalian spinal cord or hindbrain evokes locomotion. Nat Neurosci 13:246-252. Activation of Vglut2-positive cells. lumber spinal cord or caudal hindbrain is sufficient to induce locomotion.

• Kyriakatos A, Mahmood R, Ausborn J, Porres CP, Buschges A, El Manira A (2011) Initiation of locomotion in adlut zebrafish. J Neurosci 31*8422-8431.

• Kimura Y, Satou C Fujioka S, Shoji W, Umeda K, Ishizuka T, Yawo H, Higashijima S (2013) Hindbrain V2a neurons in the excitation of spinal locomotor circuits during zebrafish swimming. Curr Biol 23:1-7.

Lamprey

• Roederer E, Goldberg NH, Cohen MJ (1983) Modification of retrograde degeneration in transected spinal axons of the lamprey by applied DC current. J Neurosci 3:153-160. spinal cord transection

• Deliagina TG, Zelenin PV, Fagerstedt P, Grillner S, Orlovsky GN (2000) Activity of reticulospinal neurons during locomotion in the freely behaving lamprey. J Neurophysiol 853-863.

• Fagerstedt P, Orlovsky GN, Deliagina TG, Grillner S, Ullen F (2001) Lateral turn in the lamprey. II. Activity of reticulospinal neurons during the generation of fictive turn. J Neurophysiol 86:2257-2265.

• Brocard F, Dubuc R (2003) Differential contribution of reticulospinal cells to the control of locomotion induced by the mesencepahlic locomotor region. 90:1714-1727.

• Zelenin PV (2005) Activity of individual reticulospinal neurons during different forms of locomotion in the lamprey. Eur J Neurosci 22*2271-2282. RSのオイラー角応答性質．

• Brocard F, Dubuc R (2010) The transformation of a unilateral locomotor command into a symmetrical bilateral activation in the brainstem. J Neurosci 30:523-533.

• Derjean D, Moussaddy A, Atallah E, St-Pierre M, Auclair F, Chang S, Ren X, Zielinski B, Dubuc R (2010) A novel neural substrate for the transformation of olfactory inputs into motor output. PLoS Biol 8:e1000567.

Mouse

• Noga BR, Kriellaars DJ, Brownstone RM, Jordan LM (2003) Mechanism for activation of locomotor centers in the spinal cord by stimulation of the mesencephalic locomotor region. J Neurophysiol 90:1464-1478.

• Jordan LM, Liu J, Hedlund PB, Akay T, pearson KG (2008) Descending command systems for the initiation of locomotion in mammals. Brain Res Rev 57*183-191.

• Jordan LM () Initiation of locomotion in mammals.

• Liang H, Paxinos G, Watson C (2011) Projections from the brain to the spinal cord in the mouse. Brain Struct Funct 215:159-186.

Motor primitive/Muscle synergy

• Hart CB, Giszter SF (2010) A neural basis for motor primitives in the spinal cord. J Neurosci 30*1322-1336.

• Jing J (2009) Command systems. Encyclopedia of Neuroscience 2*1149-1158.

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その他

• Nӓssel DR, Hogmo O, Hallberg E (1984) Antennal receptors in the blowfly Calliphora erythrocephala. I. The gigantic central projection of the pedicellar campaniform sensillum. J Morphol 180:159-169. アンテナに細胞体を持つDN．唯一の下行性の感覚ニューロン．DN176

Shaw AC, Jackson AW, Holmes T, Thurman S, Daivs GR, McClellan AD (2010) Descending brain neurons in larval lamprey: Spinal projection patterns and initiation of locomotion. Exp Neurol 224:527-541.

Descending neurons in turtle

Berkowitz A, Stein PS (1994) Activity of descending propriospinal axons in the turtle hindlimb enlargement during two forms of fictive scratching: phase analysis. J Neurosci 14:5105-5119.

Flight initiation

Pearson KG, Reye DN, Parsons DW, Bicker G (1985) Flight-initating interneurons in the locust. J Neurophysiol 53:910-925.

行動を停止する下行路

Perrins R, Walford A, Roberts A (2002) Sensory activation and role of inhibitory reticulospinal neurons that stop swimming in hatchling frog tadpoles. J Neurosci 22, 4229-4240.

Li WC, Perrins R, Walford A, Roberts A (2003) The neuronal targets for GABAergic reticulospinal inhibition that stops swimming in hatchling frog tadpoles. J Comp Physiol A 189, 29-37.

Anatomy in thorocico-abdomenal ganglia

Brierley DJ, Rathore K, VijayRaghavan K, Williams DW (2012) Developmental origins and architecture of Drosophila leg motorneurons. J Comp Neurol 520:1629-1649.

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heavily invested with spines

The dendritic tree is complex in form, comprising a thickened trunk, from which arise one or several stout primary branches

CONTROL

• Zinger N, Harel R, Gabler S, Israel Z, Prut Y (2013) Functional organization of information flow in the corticospinal pathway. J Neurosci 33:1190-1197. 指とリストの動きの制御の違い．