Our central nervous system is made up of the cerebrum, the cerebellum,
the brainstem and the spinal cord. The brainstem is a trunk-like part that sits in the posterior cranial fossa and connects the spinal cord inferiorly with the forebrain superiorly. The brainstem can be divided into three parts. From caudal to rostral these parts are: the medulla oblongata, the pons and the midbrain. The brainstem is made up of white and gray matter. The white matter contains many ascending and descending fibers that act like a highway, allowing information to travel to and from the spinal cord and the higher parts of the central nervous system. Scattered amongst the white matter tracts, there are islands of gray matter that consist of neuronal cell bodies, many that are the nuclei associated with cranial nerves. Some of these collections of nuclei serve as centers for life sustaining reflexes, like those involved with breathing and our heartbeat, others coordinate states of alertness or arousal, while others mediate motor activities and relay sensory information. First, let’s look at the medulla oblongata, specifically its ventral aspect. Right in the middle, there is the anterior median fissure. On either side of it, there are two bumps called the pyramids, which contain axons of the corticospinal, or pyramidal, tract. Before entering the spinal cord, these fibers cross over to the opposite side, forming the decussation of the pyramids. Lateral to each pyramid, there are two oval bumps called the olives. They contain the inferior olivary nuclei, which have rich connections to the cerebellum and are involved in motor coordination and learning. Now, on the dorsal aspect of the medulla oblongata, it’s visible that the rostral medulla contains the inferior part of the fourth ventricle, a space filled with cerebrospinal fluid, or CSF. Across the floor of the fourth ventricle, spreading transversely, there are the striae medullaris, which are raised stripes that contain arcuatocerebellar fibers. Laterally, there are the inferior cerebellar peduncles, which contain fibers that travel between the medulla and the cerebellum. The fourth ventricle has a diamond, or rhomboid shape, so it’s also called the rhomboid fossa. The ventricle, or fossa, tapers at its caudal aspect to a point called the obex and near this caudal limit is the entrance to the central canal of the spinal cord. In this region, CSF can travel from the fourth ventricle into the central canal. Looking at the caudal medulla, right on the midline there’s the posterior median sulcus. Next to it, there are two bumps called the gracile tubercles, which contain the gracile nuclei, one on each side. Lateral to the gracile tubercles, there are two more bumps, on each side, called the cuneate tubercles, which contain the cuneate nuclei. The gracile and cuneate nuclei convey fine touch, pressure, conscious proprioception and vibratory sensations. Now, let’s switch gears and have a look at the embryological development of the medulla oblongata and the spinal cord. The development is important because it provides a location plan for motor and sensory nuclei and a blueprint for the overall pattern of motor and sensory information flow! If we look at a transverse section of the part of the neural tube that will develop into the spinal cord, we can see the cavity of the neural tube in the center. The neural tube has a dorsal, or posterior part, that forms the alar plate, and a ventral, or anterior part, that forms the basal plate. Inside the neural tube, a groove called the sulcus limitans separates these plates. Now, the alar plate gives rise to sensory nuclei and later develops into the dorsal horns of the spinal cord's gray matter. The basal plates, on the other hand, give rise to the motor nuclei and later develop into the ventral horns of the spinal cord. As we continue rostral along the neural tube to the part that will develop into the medulla oblongata, we can see that the cavity of the neural tube extends laterally to form the fourth ventricle. In doing so, it pushes the alar plates ventrally as well. Even though the basal and alar plates were ventro-dorsal in the spinal cord, they shift to a medio-lateral position in the medulla oblongata. Medially, the basal plate develops into the motor nuclei of the cranial nerves, while laterally and posteriorly the alar plate will develop into sensory nuclei. The sulcus limitans that separates them can be seen on the floor of the fourth ventricle, parallel to the posterior median sulcus. Let’s take a quick break and see if you can identify structures of the dorsal aspect of the medulla oblongata. Okay, now let’s look at the internal structures of the medulla oblongata, and the easiest way to do that is by making a few transverse sections, starting with the first one at the level of the caudal medulla. On the ventral aspect, there are the two pyramids, which contain descending fibers of the corticospinal tracts. The majority of these fibers cross over to the opposite side in the pyramidal decussation and form the lateral corticospinal tract as the fibers enter the spinal cord. Lateral to the pyramids, there’s the anterior spinocerebellar tract, and behind it, the posterior spinocerebellar tract. These carry proprioceptive input from muscle spindles, golgi tendon organs and joint receptors to the cerebellum. Medial to them, there’s the lateral spinothalamic tract, which carries somatosensory information about pain and temperature, originating from below the face, to the thalamus. Between these tracts and the decussation of the pyramids, part of the inferior olivary nucleus can be seen. On the dorsal aspect, lateral to the posterior median sulcus, the rostral part of the fasciculus gracilis can be seen on both sides and just lateral to those fibers, the fasciculus cuneatus. Ventral to them, there are the nucleus gracilis and nucleus cuneatus, which are made up of grey matter. Lateral to the fasciculus cuneatus, the spinal trigeminal tract is present and ventral to it, the spinal trigeminal nucleus. Finally, between all these nuclei and tracts, there’s the reticular formation, which is a network of nuclei with interspersed afferent and efferent fibers. It stretches across the entire brainstem, so it acts like an interface between the spinal cord and higher brain centers. The reticular formation influences many functions, such as motor reflexes, eye movements, autonomic functions and even consciousness. Now let’s look at another transverse section, this time at the level of the decussation of the medial lemnisci. Here, there’s a somatosensory tract decussation: axons originating from the nucleus gracilis and nucleus cuneatus form the internal arcuate fibers that pass around the central gray matter. Ventral to it, these fibers cross the neural axis to the opposite side. Dorsal to the decussation, in the central gray matter, there are the hypoglossal nuclei. Other structures are similar to those described in the previous section. Now, even more rostrally, let’s look at the olives and inferior cerebellar peduncles. Here, instead of the central canal dorsally, there is a space created that makes up the fourth ventricle. The central gray matter is larger and sits under the floor of the fourth ventricle, where it contains nuclei of cranial nerves, next to the posterior median sulcus. From medial to lateral, these are: the hypoglossal nuclei, dorsal motor nuclei of vagus and the solitary nucleus. Dorsal to the solitary tract, there is the medial vestibular nucleus and laterally the inferior vestibular nucleus. Ventral to all these nuclei, there’s the reticular formation, and buried deep within it the nucleus ambiguus. Lateral to the reticular formation, there are the spinal trigeminal tracts and nuclei, and most laterally, the inferior cerebellar peduncles. On the ventral side, next to the anterior median fissure, there are the pyramids. They contain the corticospinal, or pyramidal, fibers that descend to the spinal cord, and the corticobulbar fibers that target motor nuclei of the cranial nerves in the brainstem. Dorsal and lateral to the pyramids there are the inferior olivary nuclei that look like two big crumpled bags with their openings positioned towards the middle. These nuclei send fibers across the midline towards the inferior cerebellar peduncles. Also dorsal to the pyramids, and between the inferior olivary nuclei, there are the medial lemnisci, whose axons originated from the nucleus gracilis and nucleus cuneatus in the medulla. Dorsal to the medial lemnisci, and between the reticular formation, there are the tectospinal tracts that coordinate head and eye movements. Between the tectospinal tracts and the hypoglossal nuclei there are the medial longitudinal fasciculi, or MLF, which connect the vestibular and cochlear nuclei with the motor nuclei of the cranial nerves III, IV and VI. Lastly, between the reticular formation and inferior olivary nuclei are the lateral spinothalamic tracts, and laterally near the surface of the medulla are the anterior spinocerebellar tracts. Finally, let’s look at the most rostral transverse section, at the level just below the pons. Here, instead of inferior vestibular nuclei there are the lateral vestibular nuclei, positioned between the inferior cerebellar peduncles and the reticular formation. Also, dorsal to the inferior cerebellar peduncles and lateral to the fourth ventricle are the dorsal cochlear nuclei, and just ventral to those we find the ventral cochlear nuclei. Let’s take a quick break and see if you can recall the nuclei of the medulla oblongata. Let’s continue with the next part of the brainstem, which is the pons. “Pons” means bridge in latin - so the pons is like a bridge between the medulla oblongata and midbrain, ventral to the cerebellum. On its ventral aspect, there are two middle cerebellar peduncles on each side that are connected by a curved surface that looks like a bridge. In the middle of it is a shallow depression called the basilar groove, which holds the basilar artery. Switching gears, the pons also has a dorsal aspect, which is triangular shaped. It creates the rostral half of the floor of the fourth ventricle. Laterally there are the superior cerebellar peduncles. Running vertically, down the midline, stretches the median sulcus and just parallel to it, on each side, the sulcus limitans. Between the median sulcus and sulcus limitans, there are two elevations called the medial eminences, whose caudal parts enlarge to form the facial colliculi. The facial colliculi are two bumps, which are made up of axons from the facial nuclei that wrap around the abducens nuclei. Laterally, from the sulci limitans, are the vestibular areas that contain the vestibular nuclei. Caudal to the superior cerebellar peduncles we see the middle cerebellar peduncles. Now, let’s make a transverse section through the pons at the level of the facial colliculi. Dorsally, there’s the cavity of the fourth ventricle, and ventral to it, the reticular formation. Ventral to the facial colliculi are the abducens nuclei and ventro-lateral to them are the facial nuclei. The axons of the facial nuclei travel postero-medially to wrap around the abducens nuclei and then continue ventro-laterally. Lateral to the facial nuclei, we find the spinal trigeminal nucleus, and dorsal to them and lateral to the abducens nuclei are the medial vestibular nuclei. The motor tracts on this section are: the corticospinal and corticobulbar tracts, on the ventral side; and the MLF, found between the facial colliculi, around the median sulcus. As for the sensory tracts, there are: the medial lemnisci, ventral to the reticular formation; spinothalamic tracts, lateral to the reticular formation; lateral lemnisci, which carry auditory information, positioned lateral to the medial lemnisci; and the spinal trigeminal tracts, next to the spinal trigeminal nuclei. Let’s look at another transverse section, now at the level of the trigeminal nuclei. Here, we can see the motor trigeminal nuclei that sit ventral near the ventro-lateral corner of the fourth ventricle; and the main sensory trigeminal nuclei lateral to them. Other structures are similar to the previous section. Let’s take another break and see if you can identify the nuclei of the pons, as well as the important motor and sensory tracts. Finally, let’s look at the midbrain, which sits between the pons and the forebrain. First, let’s look at its ventral aspect. Laterally, there’s the crus cerebri, or the basis pedunculi, which contain descending motor tracts from the cerebrum. Between them, there’s a space called the interpeduncular fossa. The gray matter at the junction of the crus cerebri, at the base of this fossa, is called the posterior perforated substance, because many small blood vessels penetrate its surface. Switching to the dorsal aspect, we can see four bumps, or colliculi, called the corpora quadrigemina. The superior colliculi contain nuclei involved in visual reflexes, while the inferior colliculi contain nuclei that are part of the auditory pathway. Lateral to the colliculi, there are two elevations called brachia, ascending in the antero-posterior direction. The superior brachium, or brachium of the superior colliculus, stretches from the lateral geniculate body and the optic tract to the superior colliculus. It contains afferent fibers carrying information from the retina, occipital lobe and frontal eye fields of the frontal lobe. The inferior brachium, or the brachium of the inferior colliculus, stretches from the inferior colliculus to the medial geniculate body, carrying auditory information. Let’s make a transverse section through the midbrain at the level of the inferior colliculi. First, regarding its general organization, the midbrain can be divided into three main parts: the tectum, tegmentum and crus cerebri. The tectum contains both the superior and inferior colliculi and is located dorsal to the cerebral aqueduct, while the tegmentum lies ventrally. From the ventro-lateral side of the tegmentum, the crus cerebri expand, and they’re separated medially by the interpeduncular fossa. The crus cerebri together with the respective half of the tegmentum are referred to collectively as the cerebral peduncle. Around the cerebral aqueduct lies the central gray matter, or periaqueductal gray matter. Now, let’s inspect the internal structures of the midbrain. First up, the nuclei, which are the nuclei of the inferior colliculi, found dorsally, behind the central gray matter; the mesencephalic trigeminal nuclei, sitting in the central gray matter lateral to the cerebral aqueduct; the trochlear nuclei, in the ventral part of the central gray matter near the middle; and the substantia nigra, meaning black substance, found between the tegmentum and crus cerebri, separating them. The cell bodies found in the substantia nigra have granules filled with a pigment called neuromelanin, giving them a darker appearance. The substantia nigra is part of the basal ganglia, which is a system of central nervous system components that help integrate and regulate various types of information, primarily motor information. In the tegmentum, between the central gray matter and the substantia nigra, we find the reticular formation again. Moving to the motor tracts, these are: the MLF, anterior to the trochlear nucleus; and the corticospinal and corticobulbar tracts, found within the middle third of the crus cerebri. Lastly, the sensory tracts are positioned on the lateral sides of the tegmentum, behind the substantia nigra. Starting ventro-medially, these are: the medial lemnisci, the spinothalamic tracts, the trigeminal lemnisci and the lateral lemnisci. Finally, let’s move rostrally again to the level of the superior colliculi. The nuclei of the superior colliculi are posterior to the central gray matter. Lateral to them we find the pretectal nuclei that receive input from the retina about the presence of light. They send signals to the parasympathetic nuclei of the oculomotor nerve, also known as the Edinger-Westphal nuclei, situated in the ventral portion of the central gray matter. These two nuclei are involved in the pupillary light reflex. In the ventral region of the central gray matter, near the middle are the oculomotor nuclei as well. Ventro-lateral to them, in the tegmentum, there are two large nuclei called the red nuclei, which influence the tone of contraction for flexor musculature. Lateral and posterior to the red nucleus is the reticular formation. The mesencephalic trigeminal, substantia nigra, and motor and sensory tracts are present like in the previous section, but here, the lateral lemniscus is missing as it has already terminated on the nuclei of the inferior colliculus. Before the recap, let’s see if you can identify the nuclei of the midbrain, as well as the important motor and sensory tracts. Alright, as a quick recap... The brainstem is made up of the medulla oblongata, the pons and the midbrain. On the ventral aspect of the medulla there are the pyramids and the olives. Dorsally there are the inferior cerebellar peduncles, the floor of the fourth ventricle, striae medullaris, the entrance to the central canal and the gracile and cuneate tubercles. At the level of the decussation of the pyramids we can find: the central canal, pyramids, decussation of the pyramids, nuclei and fasciculi gracilis and cuneatus, spinal tracts and spinal nuclei of trigeminal, inferior olivary nuclei, ventral and dorsal spinocerebellar tracts and the lateral spinothalamic tracts. At the level of the decussation of the medial lemnisci there are the hypoglossal nuclei. At the level of the olives there are the fourth ventricle, inferior olivary nuclei, vestibular nuclei, and hypoglossal nuclei as well as the nucleus ambiguus, dorsal motor nucleus of vagus, solitary nucleuss, MLF, medial lemnisci and tectospinal tracts. Just caudal to the pons, we can see the lateral vestibular nuclei and the cochlear nuclei. On the ventral aspect of the pons we see the ventral parts of the middle cerebellar peduncles and the basilar groove. On the dorsal aspect are: the superior and middle cerebellar peduncles, median sulcus, medial eminence and facial colliculus, sulcus limitans and vestibular area. At the level of the facial colliculi, there are the facial, abducens and medial vestibular nuclei, the trigeminal lemnisci and the motor and main sensory nuclei of trigeminal, corticospinal and corticobulbar tracts, the MLF, spinothalamic tracts and medial lemnisci. The midbrain contains the crus cerebri and interpeduncular fossa on the ventral aspect and the corpora quadrigemina on the dorsal aspect along with the superior and inferior brachii and medial and lateral geniculate bodies on the lateral side. At the level of the inferior colliculi, there are: nuclei of the inferior colliculus, trochlear nuclei, substantia nigra, corticospinal and corticobulbartracts, MLF, medial lemnisci, lateral lemnisci, trigeminal lemnisci and the spinothalamic tracts. At the level of the superior colliculi there are the: nuclei of the superior colliculus, oculomotor nuclei, pretectal nuclei, Edinger-Westphal nuclei and red nuclei.