Gert Holstege

The Anatomy of Brain Stem Pathways to the Spinal Cord in Cat. A Labeled Amino Acid Tracing Study

Publisher Summary This chapter discusses the anatomy of brain stem pathways to the spinal cord in cat. In the series of experiments described in the chapter, [3H] leucine injections were made in the brain stem and the spinal distribution of the labeled descending brain stem pathways was studied autoradiographically. The pontine reticulospinal tract, as well as the vestibulospinal tracts, distribute to the ventromedial parts of the intermediate zone mainly ipsilaterally. Anterograde fiber degeneration findings obtained since the early decades of this century by means of several techniques consistently showed that the descending brain stem pathways to the spinal cord terminate mainly in the spinal intermediate zone—for example, in Rexeds laminae V–VIII. However, to facilitate the identification of the various descending pathways, the brain stem distribution of the cells of origin of these pathways in the cat has been reinvestigated by means of the horseradish peroxidase (HRP) technique by using the procedure of Mesulam.

Anatomical observations on the afferent projections to the retractor bulbi motoneuronal cell group and other pathways possibly related to the blink reflex in the cat.

In the cat retractor bulbi (RB) muscle reflexively retracts the eye ball into the orbit. This reflex action is called the nictitating membrane response which, together with the reflex contraction of the orbicularis oculi muscle, constitutes the blink reflex. The retractor bulbi (RB) motoneuronal nucleus is a small cell group located in the lateral tegmentum of the caudal pons, just dorsal to the superior olivary complex. The nucleus is identical to the accessory abducens nucleus and sends its fibers through the abducens nerve. Autoradiographical tracing results indicate that the RB nucleus receives some fibers from the principal and rostral spinal trigeminal nuclei and from the dorsal red nucleus and dorsally adjoining tegmentum. The same areas project to the intermediate facial subnucleus, containing motoneurons innervating the orbicularis oculi muscle. It is suggested that the trigeminal projections take part in the anatomical framework for the R1 component of the blink reflex. Two other brainstem areas i.e.: a portion of the caudal pontine ventrolateral tegmental field and the medullary medial tegmentum at the level of the hypoglossal nucleus were also found to project to the RB motoneuronal cell group and to the intermediate facial subnucleus. These projections were much stronger than those derived from the trigeminal nuclei and red nucleus. Moreover, the medullary premotor area projects not only to the blink motoneuronal cell groups but also to the pontine premotor area. It is suggested that both areas are involved in the R2 blink reflex component. The medullary blink premotor area receives afferents especially from oculomotor control structures in the reticular formation of the brainstem while the pontine blink premotor area receives afferents from the olivary pretectal nucleus and/or the nucleus of the optic tract and from the dorsal red nucleus and its dorsally adjoining area. Because the oculomotor control structures in the reticular formation (by way of the superior colliculus) and the red nucleus receive afferents from trigeminal nuclei, they may play an important role in tactually induced reflex blinking, while the pretectum could take part in the neuronal framework of the visually induced blink reflex.

Projections from the red nucleus and surrounding areas to the brainstem and spinal cord in the cat. An HRP and autoradiographical tracing study

HRP injections at the C2, T1 and S1 spinal levels and in the medullary lateral tegmental field revealed that the contralaterally projecting rubro-bulbospinal neurons are located not only in the caudal but also to a certain extent in the rostral red nucleus (RN). These RN projections are somatotopically organized. Neurons projecting to the sacral cord are located in the ventrolateral RN, those projecting to the upper part of the spinal cord lie in the dorsomedial RN and those projecting to the medullary lateral tegmentum were found in the dorsal portions of the RN. These last neurons are smaller than many of the other RN neurons. The HRP results also revealed that the RN does not project to the caudal raphe nuclei. The autoradiographical results confirmed the HRP findings. They further indicated that the contralateral RN projections to the caudal brainstem precerebellar nuclei (nucleus corporis pontobulbaris, lateral reticular nucleus, lateral cuneate nucleus) and the dorsal column nuclei are also somatotopically organized. This was also true for the RN projections to the dorsomedial and intermediate facial subnuclei and the caudal pontine and medullary lateral tegmental field. These areas receive afferents from mainly the dorsal portions of the RN. Regarding the RN projections to the spinal cord, the autoradiographical tracing results revealed somatotopically organized contralateral RN projections to laminae V, VI and VII. Moreover, a small but distinct RN projection to a dorsolaterally located group of motoneurons at the C8-T1 level was demonstrated. Ipsilaterally a minor projection to the cervical and upper thoracic lateral intermediate zone was observed. Finally, strong ipsilateral projections from the rostral mesencephalon to the inferior olive were seen. These projections were derived from various rostral mesencephalic areas, including the nucleus of Darkschewitsch, the nucleus accessorius medialis of Bechterew, the interstitial nucleus of Cajal and the area of the rostral interstitial nucleus of the medial longitudinal fasciculus. In the cat it was difficult to define which of the mesencephalic areas projecting to the inferior olive represented the parvocellular RN. A new subdivision of the RN is proposed based on its projections and not on the size of its cells. In this concept the first group is formed by the RN neurons projecting contralaterally to the caudal brainstem and spinal cord. The second group consists of RN neurons projecting to the inferior olive.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent projections to the orbicularis oculi motoneuronal cell group. An autoradiographical tracing study in the cat

The motoneurons innervating the orbicularis oculi muscle from a subgroup within the facial nucleus, called the intermediate facial subnucleus. This makes it possible to study afferents to these motoneurons by means of autoradiographical tracing techniques. Many different injections were made in the brainstem and diencephalon and the afferent projections to the intermediate facial subnucleus were studied. The results indicated that these afferents were derived from the following brainstem areas: the dorsal red nucleus and the mesencephalic tegmentum dorsal to it; the olivary pretectal nucleus and/or the nucleus of the optic tract; the dorsolateral pontine tegmentum (parabrachial nuclei and nucleus of Kölliker-Fuse) and principal trigeminal nucleus; the ventrolateral pontine tegmentum at the level of the motor trigeminal nucleus; the caudal medullary medial tegmentum; the lateral tegmentum at the level of the rostral pole of the hypoglossal nucleus and the ventral part of the trigeminal nucleus and the nucleus raphe pallidus and caudal raphe magnus including the adjoining medullary tegmentum. These latter projections probably belong to a general motoneuronal control system. The mesencephalic projections are mainly contralateral, the caudal pontine and upper medullary lateral tegmental projections are mainly ipsilateral and the caudal medullary projections are bilateral. It is suggested that the different afferent pathways subserve different functions of the orbicularis oculi motoneurons. Interneurons in the dorsolateral pontine and lateral medullary tegmentum may serve as relay for cortical and limbic influences on the orbicularis oculi musculature, while interneurons in the ventrolateral pontine and caudal medullary tegmentum may take part in the neuronal organization of the blink reflex.

Location of Motoneurons Innervating Soft Palate, Pharynx and Upper Esophagus. Anatomical Evidence for a Possible Swallowing Center in the Pontine Reticular Formation

Retrograde HRP studies indicated that motoneurons innervating soft palate and pharynx are located in especially the dorsal group of the nucleus ambiguus, while motoneurons innervating upper esophagus and cricothyroid muscles are located in the retrofacial nucleus, the most rostral part of the nucleus ambiguus. Autoradiographic tracing techniques revealed that in the caudal pontine tegmentum an area is located just dorsal to the superior olivary complex, that projects contralaterally to the ventral part of the trigeminal motor nucleus, the dorsal group of the nucleus ambiguus and the ventral part of the hypoglossal nucleus. In these areas motoneurons are located innervating the mylohyoid, soft palate, pharynx and geniohyoid muscles respectively. Since these muscles are all involved in swallowing, a possible involvement of this pontine tegmental area in swallowing is discussed.

Mesencephalic Projections To The Facial Nucleus In The Cat. An Autoradiographical Tracing Study

In 33 cats the projections of different parts of the mesencephalon to the facial nucleus were studied with the aid of the autoradiographical tracing method. The results indicate the existence of many different mesencephalo-facial pathways. The dorsomedial facial subnucleus, containing motoneurons innervating ear muscles, receives afferents from 4 different mesencephalic areas: a, the most rostral mesencephalic reticular formation; b, the nucleus of Darkschewitsch and/or the ventral part of the rostral PAG; c, the interstitial nucleus of Cajal and/or the mesencephalic tegmentum dorsomedial to the red nucleus. These areas project bilaterally by way of an ipsilateral medial tegmental pathway. The medial part of the deep tectum. This area projects bilaterally by way of the tecto-spinal tract. The lateral mesencephalic tegmentum close to the parabigeminal nucleus. This area projects mainly contralaterally by way of a separate contralateral lateral tegmental fiber bundle. The mesencephalic tegmentum just dorsolateral to the red nucleus and perhaps from the dorsolateral red nucleus itself. This area projects contralaterally by way of the rubrospinal tract. The intermediate facial subnucleus containing motoneurons innervating the muscle around the eye, receives afferents from two different mesencephalic areas: The dorsal part of the rostral as well as caudal red nucleus (but not from its caudal pole) and from the dorsally adjoining mesencephalic tegmentum including the area of the nucleus of Darkschewitsch and the interstitial nucleus of Cajal. These areas project contralaterally by way of the contralateral rubrospinal tract. The nucleus of the optic tract and/or the olivary pretectal nucleus. This area projects contralaterally by way of a contralateral medial tegmental pathway. The lateral and ventrolateral facial subnuclei containing motoneurons innervating the muscles around the mouth receive afferents from two different mesencephalic areas: The lateral part of the deep tectal layers. This area projects contralaterally by way of the tecto-spinal tract. The nucleus raphe dorsalis and perhaps the nucleus centralis superior. This area projects by way of the lateral tegmentum of caudal pons and medulla.

Anatomical Evidence For An Ipsilateral Rubrospinal Pathway And For Direct Rubrospinal Projections To Motoneurons In The Cat

In 11 cases [3H]leucine injections were made in various parts of the red nucleus (NR) and surrounding area. The autoradiographical tracing results confirmed the existence of a somatotopic organization in the location of the rubrospinal neurons. Moreover the results demonstrated direct NR projections to the most dorsolaterally located motoneuronal cell group in the C8 and upper T1 spinal segments and some ipsilaterally descending rubrospinal fibers terminating in the lateral part of the cervical intermediate zone.

Spinal Cord Location Of The Motoneurons Innervating The Abdominal, Cutatneous Maximus, Latissimus Dorsi And Longissimus Dorsi Muscles In The Cat.

SummaryHorseradish peroxidase (HRP) injections were made in the rectus abdominis, obliquus externus, obliquus internus, transversus abdominis, cutaneous maximus, latissimus dorsi and the longissimus dorsi muscles in the cat. The results showed that motoneurons innervating the obliquus externus, obliquus internus and transversus abdominis muscles were located in greatly overlapping areas of midthoracic, caudal thoracic and upper lumbar spinal segments. These motoneuronal cell groups were present laterally in the ventral horn and at caudal thoracic and upper lumbar levels they bordered on the white matter. The location of the rectus motoneurons differed somewhat from the location of the other motoneuronal cell groups because they were also present at low cervical and upper thoracic levels and in the segments T12 to L3 they were found in the ventral horn medial to the other abdominal muscle motoneuronal cell group. At mid-thoracic levels rectus motoneurons were located in the same area as the other abdominal muscle motoneurons. Latissimus dorsi motoneurons were observed in a large cell group in the ventrolateral part of the ventral horn at the levels caudal C6 to rostral C8. Furthermore they were found in the segments T9 to L3 laterally in the ventral horn which is the same area in which the other abdominal muscle motoneurons except the rectus ones are located. Longissimus dorsi motoneurons were located in the most ventral portion of the ventral horn in all thoracic and upper 4 lumbar segments. The cutaneous maximus motoneurons were found in a cell group, located ventrolaterally in the ventral horn at the edge of the gray and white matter at the level caudal C8-rostral T1. This cell group corresponds to the caudal part of the ventral motor nucleus (VMN) of Matsushita and Ueyama (1973). Interestingly, labeled motoneurons were also present in the VMN after injecting HRP in the abdominal muscles as well as in the caudal (but not in the rostral) parts of the latissimus dorsi muscle but not in the longissimus dorsi injected cases. The possibility whether these motoneurons are labeled because of leakage of HRP to abdominal and caudal latissimus dorsi muscles is discussed. If leakage would not be the case, motoneurons in the VMN may be involved in specific functions of the abdominal muscles, such as the so-called steady state contractions.

Differential corticospinal projections in the cat. An autoradiographic tracing study.

An autoradiographic study of the corticospinal projections from different parts of the cat sensorimotor cortex produced the following findings. The lateral part of area 4 projects contralaterally to the lateral intermediate zone of the cervical enlargement only. The intermediate part of area 4 projects throughout the spinal cord, contralaterally to the lateral part of the intermediate zone and bilaterally to its ventromedial part. The lateral and medial part of area 3 project contralaterally to the cervical and lumbosacral dorsal horn (including laminae I and II), respectively.

Anatomy Of Premotor Centers In The Reticular Formation Controlling Oculomotor, Skeletomotor And Autonomic Motor Systems

Publisher Summary This chapter discusses on the neuroanatomy of the reticular formation as a premotor relay for controlling the oculomotor, somatomotor, and autonomic motor systems. An attempt is made to inter-relate these data, and point out similarities and differences in their relation to the limbic system. The reticular formation is taken in the chapter to include a lateral, medial, and ventral region, all of which are particularly clear at pontine and medullary levels. The midline raphe nuclei are included in the medial and ventral divisions. The reticular formation has been associated with a wide variety of functions, such as arousal, eye movement generation, and posture control. The lateral zone of the reticular formation contains interneurons innervating motoneurons of the motor nuclei of the trigeminal, facial, vagal, hypoglossal and phrenic nerves, and can be regarded as the bulbar counterpart of the spinal intermediate zone. The ventral zone projects to all parts of the spinal cord throughout its length. Afferents to the ventral zone are mainly derived from limbic areas, and therefore, it may be regarded as a part of the limbic system itself. Various limbic areas also project to the lateral reticular zone, but the medial zone seems to receive no limbic afferents. This suggests that the oculomotor and postural system, unlike the somatomotor or the autonomic motor system, are not under such direct limbic control and must be considered as unique in this respect.