In memoriam: Pieter Dullemeijer (1925-2004).
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Biomedical subjects
Publications and source records attributed to J L Dubbeldam.
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This study is part of a program intended to provide the neuroanatomical framework for investigations of the role of brain areas in specific aspects of behavior in the collared dove. In the present study, the distribution of dopamine-, substance P-, vasoactive intestinal polypeptide (VIP)- and neuropeptide Y (NPY)-immunoreactivity are mapped throughout the brain of this bird. For each substance, our observations are compared with data from studies in other species of birds. Over all, our data confirm the results of previous reports, but a few differences with data from some of these studies are found. The immunohistochemical data are used in an attempt to define more precisely cell areas and their subdivisions in the avian forebrain and brainstem, and to compare these areas to nuclei in the brain of mammals.
The vestibular apparatus provides information about the position and movements of the head. Craniocervical muscles position the head with respect to the upper part of the neck. Motoneurons innervating these muscles are located in the supraspinal nucleus and ventral horn of the rostral cervical cord. Premotor neurons of craniocervical muscles have been found in the medial two-thirds of the medullary reticular formation: the ventromedial part of the parvocellular reticular formation and the gigantocellular reticular formation. In the present study, projections from vestibular nuclei upon craniocervical premotor neurons were investigated using anterograde and retrograde tracers. Vestibulospinal fibers run bilaterally in the medial vestibulospinal tract and ipsilaterally in the lateral vestibulospinal tract. Vestibuloreticular projections are mainly ipsilateral, and originate from the n. vestibularis lateralis pars ventralis and pars dorsalis, and from the n. vestibularis descendens. Terminal labeling is found in the border zone between the parvocellular and gigantocellular reticular formation. These projections show that in addition to direct bilateral vestibulo-craniocervical projections an indirect vestibular pathway to craniocervical motor nuclei exists. The direct pathway probably is the neural substrate for the vestibulocollic reflex, whereas the vestibular projection upon the reticular formation might influence head orientation during various kinds of activities, such as pecking, preening and so on.
The distribution of immunoreactivity after applying an antibody against gastrin-releasing peptide (GRP) was studied in the brain of the collared dove (Streptopelia decaocto). In the forebrain GRP-immunoreactive (GRP-ir) cells were found in the hyperstriatum accessorium, medial and lateral parts of the neostriatum, corticoidea dorsolateralis and temporoparieto-occipitalis areas, hippocampus, pre- and parahippocampal areas and prepiriform cortex. In the brainstem, GRP-ir cells were restricted mainly to the substantia nigra and ventral tegmental nucleus. Areas with densely packed GRP-ir clusters of varicosities were the medial intermediate hyperstriatum ventrale and lateral septal nucleus; dense GRP-ir neuropil was found in the parolfactory lobe, and in the dorsal half of the intermediate and caudal archistriatum. The ventral lamina medullaris contained many GRP-ir fibers. Forebrain areas devoid of immunoreactivity were the basal nucleus, ectostriatum, rostral archistriatum, most of the paleostriatum augmentatum and the lateral bed nucleus of the stria terminalis. Moderate densities of GRP-ir elements were found in the other telencephalic areas and further in, among others, the preoptic and hypothalamic region, ventral area of Tsai, cerulean nuclei, parabrachial complex, dorsal glossopharyngeal and vagus motor nuclei and medial nuclei of the solitary complex. The observations are compared with data from the literature and the implications for the definition of specific centers within the avian brain are discussed, with emphasis on systems with a role in visceral and motivational functions and in learning.
The supraspinal nucleus (SSp) in the mallard, which lies in the rostral spinal cord and caudal brainstem, is a motor nucleus that forms the rostral continuation of the ventral horn. It contains part of the motoneurons innervating the craniocervical muscles. Injections with horseradish peroxidase (HRP) and wheat germ agglutinin conjugated to HRP (WGA) in the SSp were used to localize the craniocervical premotor neurons in the medullary reticular formation. A mixture of WGA and HRP (WGA/HRP) or biotinylated dextran amine (BDA) were injected in the different reticular areas to test the results. Small numbers of craniocervical premotor neurons were found bilaterally in the ventromedial part of the parvocellular reticular formation (RPcvm) and in the caudal extension of RPcvm, the nucleus centralis dorsalis of the medulla oblongata, and the gigantocellular reticular formation (RGc). In a second series of experiments, WGA/HRP and BDA injections in these reticular areas were used to visualize afferent fibers and terminals in the SSp. The combination of the two types of experiments shows that RPcvm and RGc contain modest numbers of craniocervical premotor neurons. Because the reticular formation also contains jaw and tongue premotor neurons and receives a variety of sensory projections, the present results suggest that the medullary reticular formation plays a role in the coordination of complex movements (e.g., feeding). The pattern of afferent and efferent connections of the reticular formation is used to redefine its subdivisions in the myelencephalon of the mallard.
The distribution of four neuroactive substances was studied in the telencephalon of the collared dove using enzyme- and immunohistochemistry. The distribution of acetylcholinesterase and dopamine was similar to that described in other birds. Galanin-immunoreactive fibres were found mainly in the paleostriatum primitivum , the medial part of the parolfactory lobe (LPO) and the lateral septal nucleus; galanin may interfere with acetylcholine activity. Intense gastrin releasing peptide (GRP)-immunoreactivity was found in the neuropil of LPO, the ventral paleostriatum and the caudal archistriatum; further GRP-immunoreactive varicosities were found in the neostriatum and the hyperstriatum ventrale - particularly in its medial part - whereas GRP-immunoreactive cells occurred in the medial neostriatum, the hyperstriatum accessorium and the ventral archistriatum. These data help to define more precisely several functional telencephalic systems, but no indications for specific centers with a role in vocalization were found.
We show that a laser with a saturable absorber, described by the Yamada model, displays excitability just below threshold. A small perturbation, for example, a small input pulse, can trigger a single high output pulse, after which the system relaxes back to the off state. In order to study possible applications, such as pulse reshaping and clock recovery, approximate expressions are given for the excitability threshold and the delay between input and output pulses. Under the influence of optical noise, the system displays coherence resonance: below threshold the laser produces pulse trains with minimal jitter for a particular optimal noise level. This all-optical coherence resonance allows direct experimental verification.
The primary sensory trigeminal system in birds comprises the mesencephalic trigeminal nucleus and the trigeminal ganglion with projections to the principal sensory nucleus (PrV) and the descending tract with its subnuclei. Other cranial nerves can contribute to PrV and the descending system that together form the somatosensory system of the head. There is also a proprioceptive component. The somatosensory system comprises a component serving tactile sense and a nociceptive component. The former processes information from many mechanoreceptors in beak and tongue; both PrV and subnuclei of the descending system are involved. The nociceptive component consists of small ganglion cells projecting presumably to layers I and II of the caudal subnucleus of the descending trigeminal system and cervical dorsal horn; this is the only trigeminal region showing immunoreactivity for substance P. The effects of amputation of the tips of the beak of chickens (debeaking) are estimated by fiber counts in electron microscopic preparations of the trigeminal branches innervating that area, and by cell counts in Nissl stained sections of the trigeminal ganglion. Our data indicate that debeaking causes a loss of exteroceptive units, but not of nociceptive units. Comparison of sections stained for the presence of substance P (immunohistochemistry) did not reveal a long-term effect on the nociceptive system suggestive of the occurrence of chronic pain.
The optic tectum in birds receives visual information from the contralateral retina. This information is passed through to other brain areas via the deep layers of the optic tectum. In the present study the crossed tectobulbar pathway is described in detail. This pathway forms the connection between the optic tectum and the premotor area of craniocervical muscles in the contralateral paramedian reticular formation. It originates predominantly from neurons in the ventromedial part of stratum griseum centrale and to a lesser extent from stratum album centrale. The fibers leave the tectum as a horizontal fiber bundle, and cross the midline through the caudal radix oculomotorius and rostral nucleus oculomotorius. On the contralateral side fibers turn to ventral and descend caudally in the contralateral paramedian reticular formation to the level of the obex. Labeled terminals are found in the ipsilateral medial mesencephalic reticular formation lateral to the radix and motor nucleus of the oculomotor nerve, and in the contralateral paramedian reticular formation, along the descending tract. Neurons in the medial mesencephalic reticular formation in turn project to the paramedian reticular formation. Through the crossed tectobulbar pathway visual information can influence the activity of craniocervical muscles via reticular premotor neurons.
The reticular formation of the brainstem contains premotor systems for various musculomotor systems. In this paper, the bulbar premotor systems for jaw and tongue movements, head and neck movements, locomotion, and respiration and vocalization in birds are reviewed and compared to premotor systems in mammals. Roughly, the bulbar reticular formation can be subdivided in three longitudinal zones: a dorsolateral (RPcdl) and a ventromedial (RPcvm) parvocellular zone and a gigantocellular zone (RGc). RPcdl contains premotor neurons for the jaw and neck system, RPcvm for the jaw, tongue and neck system, and RGc for the tongue and locomotory system. RPcdl receives input from the descending sensory trigeminal system, parts of RPcvm and RGc from vestibular nuclei, whereas the tectum has a projection to the contralateral RGc. RPcdl and RPcvm receive substantial telencephalic input through the occipitomesencephalic tract. The bulbar part of the respiratory system consists of a series of cell groups in the ventrolateral reticular formation and has connections with motor centers of the vocalization system. The similarities and differences between the avian and mammalian situation are discussed. Musculomotor systems participate in various activities. It is argued that a premotor system should possess sufficient flexibility to control the participation of a motor system in the different activities. This flexibility may permit the occurrence of learning processes in terms of refining basically existing motor patterns. The emergence of new and more complex motor patterns as in vocalization requires the involvement of hierarchically higher brain centers.
The intratelencephalic and descending connections of the archistriatum of the mallard were studied using anterograde and retrograde tracers. Autoradiography after injections of [3H]-leucine served to visualize the intratelencephalic and extratelencephalic efferent connections of the archistriatum. Horseradish peroxidase (HRP), HRP-wheatgerm agglutinin, and fluorescent tracers were used to identify the precise origin of the projections to the various terminal fields found in the anterograde experiments. Four main regions can be recognized in the archistriatum of the mallard: (1) the rostral or anterior part that is a source of contralateral intratelencephalic projections, in particular to the contralateral archistriatum; (2) the dorsal intermediate archistriatum that is the origin of a large descending fiber system, the occipitomesencephalic tract, with projections to dorsal thalamic nuclei, the medial spiriform nucleus, the intercollicular nucleus, the deep tectum, parts of the mesencephalic and bulbar reticular formation, and the subnuclei of the descending trigeminal tract. There are no direct projections to motor nuclei. This part corresponds to the somatic sensorimotor part as defined by Zeier and Karten (1971, Brain Res. 31:313-326); it also contributes to the ipsilateral intratelencephalic connections and, to a lesser degree, to contralateral intratelencephalic connections. (3) The ventral intermediate archistriatum is another region that is also a source of intratelencephalic projections, in particular of those to the lobus parolfactorius. The most lateral zone sends fibers to the septal area. (4) The caudoventral intermediate and posterior archistriatum is another region that is a source of the projections to the hypothalamus and thus corresponds to the amygdaloid part of the archistriatum as defined by Zeier and Karten; it also contributes a modest component to the occipitomesencephalic tract. The different cell populations are not spatially separated, which makes it impossible to recognize distinct subnuclei within the four main regions of the archistriatum of the mallard.
BACKGROUND: In the mallard duck, functionally distinct groups of jaw muscles are each innervated by a different subnucleus of the main trigeminal (mV) or facial (mVII) motor nucleus. The other subnuclei of mV and mVII innervate several head muscles, including lingual muscles. The reticular premotor cells of the trigeminal and facial jaw motor subnuclei occupy different areas in the parvocellular reticular formation (RPc). The cell bodies of jaw muscle spindle afferents are situated in the mesencephalic nucleus (MesV). In the present study, the central connections of MesV with jaw motor subnuclei and their premotor areas are investigated. METHODS: In a first series of experiments, horseradish peroxidase (HRP) injections were made in electrophysiologically identified trigeminal and facial subnuclei. In a second series of experiments, HRP was delivered iontophoretically at different parts of RPc. Anterograde tracing with tritiated leucine was used to confirm the central connections of MesV. Double labeling with fluorescent tracers was used to investigate whether MesV collaterals reach both the rostral and caudal parts of RPc. RESULTS: MesV projects to only two of the five different subnuclei of the trigeminal motor nucleus. The subnuclei that receive spindle afferents innervate jaw adductor muscles (mV2) or pro- and retractors of the mandible (pterygoid muscles; mV1). The three other subnuclei innervate jaw-opener muscles or other head muscles. MesV fibers also project to the rostral part of the dorsolateral RPc (RPcdl), which serves as a premotor area for the motor subnuclei of adductor and pterygoid muscles. The intermediate part of RPcdl does not contain premotor cells of mV or mVII, and a clear projection of MesV to this area is absent. The caudal part of RPcdl projects to the mV and mVII subnuclei that innervate jaw-opener muscles. This part of RPc receives a projection from the same MesV cells as the rostral RPcdl. The MesV projection to RPc does not include premotor cells of mV and mVII in the ventromedial part of RPc (RPcvm). CONCLUSIONS: Spindle afferents from jaw-closer muscles project only to mV subnuclei innervating jaw-closer muscles (mV1, mV2) and to a population of premotor cells in the rostral RPcdl that innervates these subnuclei. The mixed population of premotor cells in RPcvm, which innervates both jaw-opener and jaw-closer subnuclei, does not receive a MesV projection. However, a premotor area for jaw-opener subnuclei in the caudal part of RPcdl does receive MesV input and may serve as a relay through which proprioceptive information from jaw closer spindles can reach jaw opener muscles.
As part of a study concerning the organization of premotor areas in the medullary reticular formation in birds we used a fluorescent retrograde double labeling technique to localize the premotor neurons of the trigeminal (mV) and supraspinal motor nucleus (SSp). Diamidino Yellow injections in mV and Fast Blue injections in SSp demonstrated that mV and SSp do not share premotor neurons, but the premotor neurons form a mixed population in the ventromedial part of the parvocellular reticular formation.
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The long term effects of amputation of the tip of the beak were studied in adult hens that were debeaked on the day of hatching, at the age of 8 d and at 6 wk, by EM analysis of fibre spectra of the medial branch of the ophthalmic nerve and of the intramandibular nerve. Three categories of fibre were distinguished for further analysis, i.e. unmyelinated axons, small myelinated fibres and large myelinated fibres. In normal birds the ophthalmic nerve contains relatively more large fibres than the intramandibular nerve. Amputation consistently results in a reduction of the number of large fibres and a substantial increase in the number of small myelinated fibres. The proportion of unmyelinated axons is rather variable, but is not affected by beak trimming. Age at debeaking has no effect. The observations are inconclusive concerning the possibility of heightened nociception.
Two sensorimotor 'feeding' circuits and their descending projections are described in the mallard and compared to those in the pigeon. The tactile/trigeminal circuit consists of a pattern of reciprocal connections of the nucleus basalis with the overlying parts of the frontal neostriatum (Nf) and hyperstriatum ventrale (HV). The dorsal zone of Nf projects to the sensorimotor part of archistriatum, to the paleostriatum augmentatum (PA) and to the lateral lobus parolfactorius. A comparable pattern of visual connections has been found: the ectostriatum has reciprocal connections with the intermediate neostriatum (Ni) and HV. Here, too, Ni has projections to archistriatum and probably to PA. Archistriatum is the source of a large descending pathway with, among others, substantial projections to the parvocellular reticular formation of the brain stem. This provides a pathway for telencephalic control of premotor systems of jaw, tongue and neck muscles. The paleostriatal complex is the source of the ansa lenticularis with projections to the nucleus spiriformis lateralis and the nucleus tegmenti pedunculopontinus, both projecting to the deep tectum. This pathway may be important for the control of body position. In the mallard, it includes a substantial trigeminal component that has not been found in the pigeon.
The trigeminal and facial motor nuclei in the mallard comprise several subnuclei which innervate tongue, jaw and other head muscles. The premotor cells of the subnuclei innervating jaw muscles are distributed in two longitudinal cell columns within the parvocellular reticular formation (RPc). The ventromedial part of RPc contains cells projecting to subnuclei innervating either jaw-closer muscles or jaw-opener muscles. In the dorsolateral part of RPc three subdivisions may be recognized: a rostral part which projects to two trigeminal subnuclei innervating jaw-closer muscles, an intermediate part which does not serve as premotor area for any of the jaw motor subnuclei and a caudal part of RPcdl which projects to the trigeminal and facial motor subnuclei innervating jaw-opener muscles. Exteroceptive information from mechanoreceptors in the beak reaches all three parts of RPCdl. Muscle spindles in jaw-closer muscles may influence the activity of premotor cells of jaw-opener muscles through their projection upon the caudal part of RPcdl.
Reticular premotor neurons of craniocervical muscles in the duck were localized with the retrograde tracer HRP and the anterograde tracer WGA-HRP. In the reticular formation neck premotor neurons were found in the gigantocellular reticular nucleus and in the ventromedial part of the parvocellular reticular nucleus rostral to the obex, and caudal to the obex in the nucleus centralis ventralis of the medulla. Results were compared with premotor areas of jaw muscles. The ventromedial part of the parvocellular reticular formation contains neck as well as jaw premotor neurons. This area may serve as the neural substrate for the coordination of neck and jaw movements.