The circuit from the cochlear nucleus magnocellularis to the nucleus laminaris supports the encoding and measurement of interaural time differences in the auditory brainstem. Specializations for the encoding of temporal information include the few and/or short dendrites and thick axons of the magnocellular and laminaris neurons, and the high degree of convergence in the circuit. Magnocellular cells have large cell bodies covered with somatic spines. The cells have few dendrites, and the number of dendrites decreases from low to high best frequency regions of the nucleus. Magnocellular neurons receive both auditory nerve terminals and GABAergic terminals with symmetric synapses and terminals filled with pleomorphic vesicles. The axonal projections of magnocellular neurons to the nucleus laminaris form maps of interaural time difference. About 100 magnocellular afferents from each side converge on each laminaris neuron, and the terminals from each side do not occupy separate domains on the cell. These terminals form punctate asymmetric synapses on both the dendrites and the cell bodies of laminaris neurons. Laminaris neurons also receive GABAergic terminals which form symmetric synapses. Laminaris neurons have oval cell bodies covered with very short dendrites. The cells in the low best frequency region of the nucleus laminaris have longer dendrites.
The central projections of the auditory nerve were examined in the barn owl. Each auditory nerve fiber enters the brain and divides to terminate in both the cochlear nucleus angularis and the cochlear nucleus magnocellularis. This division parallels a functional division into intensity and time coding in the auditory system. The lateral branch of the auditory nerve innervates the nucleus angularis and gives rise to a major and a minor terminal field. The terminals range in size and shape from small boutons to large irregular boutons with thorn-like appendages. The medial branch of the auditory nerve conveys phase information to the cells of the nucleus magnocellularis via large axosomatic endings or end bulbs of Held. Each medial branch divides to form 3-6 end bulbs along the rostrocaudal orientation of a single tonotopic band, and each magnocellular neuron receives 1-4 end bulbs. The end bulb envelops the postsynaptic cell body and forms large numbers of synapses. The auditory nerve profiles contain round clear vesicles and form punctate asymmetric synapses on both somatic spines and the cell body.
The barn owl's head grows after hatching, causing interaural distances to more than double in the first 3 weeks posthatch. These changes expose the bird to a constantly increasing range of interaural time cues. We have used Golgi and ultrastructural techniques to analyze the development of the connections and cell types of the nucleus magnocellularis (NM) and the nucleus laminaris (NL) with reference to the growth of the head. The time coding circuit is formed but immature at the time of hatching. In the month posthatch, the auditory nerve projection to the NM matures, and appears adult-like by posthatch day (P)21. NM neurons show a late growth of permanent dendrites starting at P6. Over the first month, these dendrites change in length and number, depending upon rostrocaudal position, to establish the adult pattern in which high best frequency neurons have few or no dendrites. These changes are not complete by P21, when NM neurons still have more dendrites than in the adult owl. The neurons of NL have many short dendrites before hatching. Their number is greatly reduced by P6, and then does not change during later development. Like NM neurons, NL neurons and dendrites grow in the first month posthatch, and at P21, NL dendrites are longer than those in the adult owl. Thus, the auditory brainstem circuits grow in the first month after hatching, but are not yet mature at the time the head reaches its adult size.
Ligules of Selaginella pilifera and S. uncinata were studied by light and electron microscopy. These ligules can be anatomically divided into tip, neck, and basal regions. The upper part of the ligule base is composed of very dense polygonal cells containing many ribosomes, endoplasmic reticulum (ER), and Golgi bodies. In S. pilifera, callose-like walls are secreted around these cells. The lower part of the base consists of wedge-shaped glossopodial cells and the adjacent two layers of sheath cells. Transfer-cell-like walls separate the glossopodium and upper sheath layer, whereas walls containing prominent plasmodesmata separate the two sheath layers. A continuous cuticle covers the entire ligule. During early ontogeny, all ligule cells are highly RNA positive, whereas later only the polygonal base cells stain densely. The anatomy and development of the ligule suggest that it is an active structure. The results of this study are compared with previous studies, especially in relation to ligular function.
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