Добавил:
Sekretar
kiopkiopkiop18@yandex.ru
t.me/Prokururor I Вовсе не секретарь, но почту проверяю
Опубликованный материал нарушает ваши авторские права? Сообщите нам.
Вуз:
Предмет:
Файл:Ординатура / Хирургия / Библиотека им академика М.И. Перельмана / Книга_4433_Библиотеки_им_академика_М_И_Перельмана
.pdf
34 BALANCE FUNCTION ASSESSMENT AND MANAGEMENT
https://t.me/medicina_free
that chicks are able to walk and run within hours of
hatching, show that in the chick the vestibular periphery matures early and is functional even before hatch.
DEVELOPMENTAL MILESTONES OF
VESTIBULAR
FUNCTION AND BALANCE
BEHAVIORS IN THE HUMAN
By the twelfth to fourteenth weeks of gestation, the
human vestibular epithelium appears almost mature
and contains types I and II hair cells, myelinated calyxbearing and bouton-terminated afferent neurons, as
well as efferent axon terminals (Dechesne, 1992). Ten
weeks later (week 24) they are considered adult-like,
and at birth, some 10 to 16 weeks later (weeks 36 to
40), the periphery is considered mature. Although
the vestibular periphery may be mature at birth in
the human, this is not readily apparent from human
behavior (unlike the case in the chick above). The
human neonate requires years (see below) to acquire
a mature vestibular-linked behavioral repertoire. It is
clear that central myelination and circuit refinements
must take place in order to fully equip the otherwise
largely immature cortical, cerebellar, extrapyramidal,
and brainstem descending motor control systems for
the support of mature behavioral motor programs and
postures. Behaviors such as walking and running as
well as mature posturing must develop after birth in
the human (e.g., sitting up, rolling over, even holding
one’s head up; see Table 2–2; Eviatar & Eviatar, 1978;
Frankenburg & Dodds, 1967). Table 2–2 summarizes
well-known developmental milestones for motor skills
in the human infant. Central immaturities therefore can
obscure or mask true vestibular functional capability as
well as deficits in neonates and children. Moreover, failure to recognize vestibular deficits may, in the absence
of intervention, put individuals at risk for developing
abnormal postural, dynamic motor control and/or
motor effector systems (e.g., De Kegel et al., 2012; Fife
et al., 2000; Shall, 2009; Van Cleave & Shall, 2006)
A functional vestibular periphery is evident in
the human neonate. Full-term normal infants typically
demonstrate vestibular mediated ocular compensation
induced by head rotation (Cyr et al., 1985; Donat et al.,
1980; Eviatar & Eviatar, 1978; Eviatar et al., 1974, 1979;
Ornitz et al., 1979; Staller, 1986; Tibbling, 1969; WienerVacher et al., 1996). Head rotation elicits a bilateral
response from the vestibular periphery. VOR measurements at the youngest ages generally show tonic conjugate deviation of the eyes in a direction opposite that
of rotation, and may include occasional rapid saccades.
The slow compensatory movement is dependent on
the vestibular periphery, whereas saccades depend on
brainstem circuitry (see Chapters 3 and 4). A reduced
number or frequency of saccades reflects the immature
state of brainstem neural circuits. The presence of saccades in the neonate is in part dependent on frequency
of rotation. Saccades may be absent in very young or
premature infants but are found consistently in normal
1- to 3-month-old babies.
Caloric testing in the neonate evaluates each ear
separately and has often provided evidence of central
immaturity, especially in premature infants (Eviatar
et al., 1974, 1979; Donat et al., 1980). VOR responses
to calorics are variable and may be absent in infants
less than 6 months. By 3 to 6 months a consistent VOR
is generally present (Cyr et al., 1985, Eviatar et al.,
1974, 1979; Donat et al., 1980). The absence of a VOR at
Table 2–2. Familiar Developmental Milestones for Motor Skills in the Human Infant*
Motor Skill 25% (mo) 50% (mo) 75% (mo) 90% (mo)
Lifts head (prone) 0.7
Lifts head 90 degrees (prone) 1.3 2.2 2.6 3.2
Chest up, arm support (prone) 2.0 3.0 3.5 4.3
Sits (head steady) 4.8 5.5 6.5 7.8
Rolls over 2.3 2.8 3.8 4.7
Stands holding on 5.0 5.8 8.5 10.0
Walks well 11.3 12.1 13.5 14.3
*Selected data from Frankenberg and Dodds (1967). Age (months) at which the respective fraction
(25%, 50%, 75%, and 90%) of infants successfully completed motor tasks listed.
2. ontogEny oF tHE vEstiBulAr systEm And BAlAnCE 35
https://t.me/medicina_free
10 months or older is considered abnormal (Fife et al.,
2000). Most abnormal responses (e.g., disconjugate eye
movements, tonic ocular deviation, absent saccades)
were found in premature or very young normal term
babies (Donat et al., 1980; Tibbling, 1969), but such
abnormalities generally reflected central immaturities
which disappeared with maturation. Thus, improvements in vestibular responses largely represent central
maturational processes which occur over several years.
Vestibular reflex testing and posturography also
have been used to assess the time course of maturation
in children and demonstrate that the process begins
very early in the neonate and continues into adolescence. There is considerable variation in the VOR findings for some parameters in growing children. Results
depend on the particular metric used to characterize
eye movements and the range of subject ages represented. Measurements can also be affected by the
subjects’ state of alertness. The ability to arouse and
maintain alertness throughout testing is itself a function of age and this can influence results.
Using Barany rotary chair testing, a number of
investigators have reported a decrease in velocity of the
VOR slow component with maturation (Ornitz et al.,
1979; Tibbling, 1969; Wiener-Vacher et al., 1996). Maturational changes appeared to be larger during the first
year after birth and change more slowly thereafter into
adolescence. Although these results outline a temporal
profile, it is difficult to relate the findings to specific
components of the vestibular ocular reflex.
VOR gain provides better insight regarding the
effectiveness of vestibular reflex compensation for
head motion. VOR gain reportedly increased linearly
in a group of 120 children studied longitudinally over
the period from 3 to 9 years of age (e.g., mean gains,
3 years: 0.58 to 9 years: 0.82, for 0.5 Hz rotations; Casselbrant et al., 2010), whereas thereafter from young
adolescence to the adult, gain reportedly may decrease
slightly at some frequencies (Herman et al., 1982;
Valente, 2007). The findings of Charpiot et al. (2010)
also indicated a decreasing gain for a slightly older
group of children (6 to 12 years old). Others found
little difference in VOR gain over ages studied (7 to
12 years old; Horak et al., 1988; 3 months to 6 years:
Cyr et al., 1985). However, the range of ages and frequencies studied were somewhat different and sample
sizes were smaller in the latter cases, which may in part
account for differences in findings. At any given age the
variance for VOR gain in humans is quite large (e.g.,
Peterka et al., 1990a, 1990b). Nonetheless, there is support for the hypothesis that a slow systematic maturation of VOR gain (increasing mean) occurs in the young
child through preadolescence and small reductions in
mean gain from the second to third decade of life. In
addition to that cited above, evidence for this comes
from a large cross-sectional study of 261 individuals,
ages 7 to 81 years, which indicates a similar temporal
profile where VOR gains appear highest for ages 7 to
20 years and then decrease somewhat and remain relatively stable over the ages from 30 to 81 years (Peterka
et al., 1990a, 1990b). Changes in VOR gain are small
relative to the variance in any case. Controversy persists in the literature regarding maturation over ages
from approximately 10 to 30 years. In the future, it will
be important to use standardized methods and metrics
including standard frequencies and rotational velocities to minimize variability across studies. It would be
helpful to see large longitudinal studies providing normative data for VOR in children and adults through the
third decade of life. Such studies, although difficult to
achieve, would serve to clarify the profile of change in
the VOR over the human lifetime.
Computerized dynamic posturography (CDP)
includes a sensory organization test (SOT), which provides a measure of how effectively sensory information is used by an individual to maintain balance and
stability under dynamic postural challenge (see Chapter 15). SOT examines how one uses visual, somatosensory, and vestibular information to maintain balance
and quantifies, among other things, the amount of
sway exhibited. Several investigators have used the
SOT in children between the ages of 3 and 15 years to
characterize changes in balance skills with maturation
and to develop normative data on sensory weighting strategies (e.g., Casselbrant et al., 2010; Charpiot,
Tringali, Ionescu, Vital-Durand, & Ferber-Viart, 2010;
Hirabayashi & Iwasaki, 1995; Peterka & Black, 1990;
Peterson et al., 2006; Valente, 2007). The evidence suggests a steady improvement on vestibular tasks that
continues into late childhood and by some reports
into adolescent years when scores approached those of
adults. In general, skills in the use of vestibular sensory
input were the last to mature relative to use of somatosensory and visual cues for maintaining balance.
In summary, whereas the peripheral vestibular
apparatus is essentially mature at birth, central maturation continues into adolescence. The VOR can be
elicited in the neonate and it matures substantially
over the first three years. Improvement generally continues into adolescence when adult balance skills are
achieved. Although progress in our understanding has
been made, it should be clear that we are only meagerly
informed about the complex multisensory process of
maturation in human postural balance and vestibular reflex systems. Our understanding of human vestibular development would benefit substantially from
36 BAlAnCE FunCtion AssEssmEnt And mAnAgEmEnt
https://t.me/medicina_free
additional research aimed at establishing normative
data over wider age ranges and improving our ability
to make functional assessments at all ages.
referenCes
Anniko, M. (1983a). Embryonic development of vestibular
sense organs and their innervation. In R. Romand (Ed.),
Development of auditory and vestibular systems (pp. 375–423).
New York, NY: Guilford Press.
Anniko, M. (1983b). Cytodifferentiation of cochlear hair cells.
American Journal of Otolaryngology, 4, 375–388.
Balaban, C. D. (2002). Neural substrates linking balance con-
trol and anxiety. Physiology & Behavior, 77, 469–475.
Balaban, C. D., & Porter, J. D. (1998). Neuroanatomic sub-
strates for vestibulo-autonomic interactions. Journal of
Vestibular Research, 8, 7–16.
Bermingham, N. A., Hassan, B. A., Price, S. D., Vollrath, M.
A., Ben-Arie, N., Eatock, R. A., . . . Zoghbi, H. Y. (1999).
Math1: An essential gene for the generation of inner ear
hair cells. Science, 284, 1837–1841.
Beurg, M., Fettiplace, R., Nam, J.-H., & Ricci, A. J. (2009).
Localization of inner hair cell mechanotransducer channels using high-speed calcium imaging. Nature Neurosci-
ence, 12, 553–558.
Bok, J., Chang, W., & Wu, D. K. (2007). Patterning and mor-
phogenesis of the vertebrate inner ear. International Journal
of Developmental Biology, 51, 521–533.
Brandt, N., Kuhn, S. Münker, S., Braig, C., Winter, H., Blin,
N., . . . Engel, J. (2007). Thyroid hormone deficiency affects
postnatal spiking activity and expression of Ca
channels in rodent inner hair cells. Journal of Neuroscience,
27(12), 3174–3186.
Bredberg, G. (1968). Cellular pattern and nerve supply of the
human organ of Corti [Supplemental material]. Acta Oto-
Laryngologica, 236, 1–135.
Bruce, L. L., Christensen, M. A., & Warr, B. (2000). Postnatal
development of efferent synapses in the rat cochlea. Jour-
nal of Comparative Neurology, 423, 532–548.
Bruce, L. L., Kingsley, J., Nichols, D. H., & Fritzsch, B. (1997).
The development of vestibulocochlear efferents and
cochlear afferents in mice. International Journal of Develop-
mental Neuroscience, 15(4–5), 671–692.
Bryant, J. E., Forge, A., & Richardson, G. P. (2005). The dif-
ferentiation of hair cells. In M. W. Kelley, D. Wu, A. A.
Popper, & R. R. Fay (Eds.), Development of the inner ear (pp.
158–203). New York, NY: Springer-Verlag.
Casselbrant, M. L., Mandel, E. M., Sparto, P. J., Perera, S., Red-
fern, M. S., Fall, P.A., . . . Furman, J. (2010). Longitudinal
posturography and rotational testing in children three to
nine years of age: Normative data. Otolaryngology-Head
and Neck Surgery, 142, 708–714.
Chambard, J. M., Chabbert, C., Sans, A., & Desmadryl, G.
(1999). Developmental changes in low and high voltage-
2+
and K+
activated calcium currents in acutely isolated mouse vestibular neurons. Journal of Physiology, 518, 141–149.
Charpiot, A., Tringali, S., Ionescu, E., Vital-Durand, F., & Fer-
ber-Viart, C. (2010). Vestibulo-ocular reflex and balance
maturation in healthy children aged from six to twelve
years. Audiology & Neurotology, 15, 203–210.
Chen, J. W. Y., & Eatock, R. A. (2000). Major potassium con-
ductance in type I hair cells from rat semicircular canals:
Characterization and modulation by nitric oxide. Journal
of Neurophysiology, 84, 139–151.
Chen, P., Johnson, J. E., Zoghbi, H. Y., & Segil, N. (2002). The
role of Math1 in inner ear development: Uncoupling the
establishment of the sensory primordium from hair cell
fate determination. Development, 129, 2495–2505.
Chen, P., & Segil, N. (1999). P27kip1 links cell proliferation
to morphogenesis in the organ of Corti. Development, 126,
1581–1590.
Cioni, G., Favilla, M., Ghelarducci, B., & La Noce, A. (1984).
Development of the dynamic characteristics of the horizontal vestibulo-ocular reflex in infancy. Neuropediatrics,
15(3), 125–130.
Correia, M. J., Rennie, K. J., & Koo, P. (2001). Return of potas-
sium ion channels in regenerated hair cells. Possible
pathways and the role of intracellular calcium signaling. Annals of the New York Academy of Sciences, 942, 228–
240.
Curthoys, I. S. (1983). Development of function of primary
vestibular neurons. In R. Romand (Ed.), Development of
auditory and vestibular systems (pp. 425–461). New York,
NY: Academic Press.
Cyr, D. G., Brookhouser, P. E., Valente, M., & Grossman, A.
(1985). Vestibular evaluation of infants and preschool children. Journal of Otolaryngology-Head and Neck Surgery, 93,
463–468.
Davies, A. M. (1996). The neurotrophic hypothesis: Where
does it stand? Philosophical Transactions of the Royal Society
of London B, 351, 389–394.
Dechesne, C. J. (1992). The development of vestibular sen-
sory organs in human. In R. Romand (Ed.), Development of
auditory and vestibular systems 2 (pp. 419–447). Amsterdam,
Netherlands: Elsevier.
Dechesne, C. J., Rabejac, D., & Desmadryl, G. (1994). Devel-
opment of calretinin immunoreactivity in the mouse inner
ear. Journal of Comparative Neurology, 346, 517–529.
De Kegel, A., Maes, L., Baetens, T., Dhooge, I., & Van
Waelvelde H. (2012). The influence of a vestibular dysfunction on the motor development of hearing-impaired
children. Laryngoscope, 122, 2837–2843.
Denman-Johnson, K., & Forge, A. (1999). Establishment of
hair bundle polarity and orientation in the developing
vestibular system of the mouse. Journal of Neurocytology,
28, 821–835.
Desmadryl, G., Dechesne, C., & Raymond, J. (1992). Recent
aspects of development of the vestibular sense organs
and their innervation. In R. Romand (Ed.), Development of
auditory and vestibular systems 2 (pp. 461–487). Amsterdam,
Netherlands: Elsevier.
2. ontogEny oF tHE vEstiBulAr systEm And BAlAnCE 37
https://t.me/medicina_free
Desmadryl, G., Raymond, J., & Sans, A. (1986). In vitro elec-
trophysiological study of spontaneous activity in neonatal
mouse vestibular ganglion neurons during development.
Developmental Brain Research, 25, 133–136.
Donat, J. F. G., Donat, J. R., & Lay, K. S. (1980). Changing
response to caloric stimulation with gestational age in
infants. Neurology, 30, 776–778.
Driver, E. C., Sillers, L., Coate, T. M., Rose, M. F., & Kelley, M.
W. (2013). The Atoh1-lineage gives rise to hair cells and
supporting cells within the mammalian cochlea. Develop-
mental Biology, 376, 86–98.
Eatock, R. A., & Hurley, K. M. (2003). Functional develop-
ment of hair cells. In R. Romand & I. Vareal-Nieto (Eds.),
Development of auditory and vestibular systems 3 (pp. 389–
448). Amsterdam, Netherlands: Elsevier.
Eatock, R. A., & Rüsch, A. (1997). Developmental changes in
the physiology of hair cells. Seminars in Cell & Developmen-
tal Biology, 8, 265–275.
Eatock, R. A., Xue, J., & Kalluri, R. (2008). Ion channels in
mammalian vestibular afferents may set regularity of firing. Journal of Experimental Biology, 211, 1764–1774.
Ernfors, P., Van de Water, T., Loring, J., & Jaenisch, R. (1995).
Complementary roles of BDNF and NT-3 in vestibular and
auditory development. Neuron, 14, 1153–1164.
Eviatar, L., & Eviatar, A. (1978). Neurovestibular examination
of infants and children. Advanced Oto-Rhino-Laryngology,
23, 169–191.
Eviatar, L., & Eviatar, A. (1979). The normal nystagmic
response of infants to caloric and perrotatory stimulation.
Laryngoscope, 89, 1036–1045.
Eviatar, L., Eviatar, A., & Naray, I. (1974). Maturation of neu-
rovestibular responses in infants. Developmental Medicine
and Child Neurology, 16, 435–446.
Faulstich, B. M., Onori, K. A., & du Lac, S. (2004). Comparison
of plasticity and development of mouse optokinetic and
vestibulo-ocular reflexes suggests differential gain control
mechanisms. Vision Research, 44, 3419–3427.
Favre, D., & Sans, A. (1978). The development of vestibular
efferent nerve endings during cat maturation: Ultrastructural study. Brain Research, 142, 333–337.
Fekete, D. M., & Camparo, A. M. (2007). Axon guidance in
the inner ear. International Journal of Developmental Biology,
51, 549–556.
Fife, T. D., Tusa, R. J., & Furman, J. M. (2000). Assessment:
Vestibular testing techniques in adults and children:
Report of the Therapeutics and Technology Assessment
Subcommittee of the American Academy of Neurology.
Neurology, 55, 1431–1441.
Forge, A., Souter, M., & Denman-Johnson, K. (1997). Struc-
tural development of sensory cells in the ear. Seminars in
Cell & Developmental Biology, 8, 225–237.
Frankenburg, W. K., & Dodds, J. B. (1967). The Denver Devel-
opmental Screening test. Journal of Pediatrics, 71, 181–191.
Fritzsch, B., Beisel, K. W., Jones, K., Farinas, I., Maklad, A.,
Lee, J., & Reichardt, L. F. (2002). Development and evolution of inner ear sensory epithelia and their innervations.
Journal of Neurobiology, 53, 143–156.
Fritzsch, B., & Nichols, D. H. (1993). DiI reveals a prenatal
arrival of efferents at the differentiating otocyst of mice.
Hearing Research, 65, 51–60.
Fritzsch, B., Silas-Santiago, I., Bianchi, L. M., & Farinas, I.
(1997). The role of neurotrophic factors in regulating the
development of inner ear innervation. Trends in Neurosci-
ence, 20(4), 159–164.
Fuchs, P. A., & Sokolowski, B. H. A. (1990). The acquisition
during development of Ca++ activated potassium currents by cochlear hair cells of the chick. Proceedings of the
Royal Society of London B, 241, 122–126.
Galicia, S., Cotes, C., & Galindo, F. (2010) Development of
spontaneous activity and response properties of primary
lagenar neurons in the chick. Cell and Molecular Neurobiol-
ogy, 30, 327–331.
Géléoc, G. S. G., & Holt, J. R. (2003). Developmental acqui-
sition of sensory transduction in hair cells of the mouse
inner ear. Nature Neuroscience, 6(10), 1019–1020.
Géléoc, G. S. G., Risner, J. R., & Holt, J. R. (2004). Develop-
mental acquisition of voltage-dependent conductances
and sensory signaling in hair cells of the embryonic mouse
inner ear. Journal of Neuroscience, 24(49), 11148–11159.
Ginzberg, R. D., & Gilula, N. B. (1979). Modulation of cell
junctions during differentiation of the chicken otocyst sensory epithelium. Developmental Biology, 68, 110–129.
Groves, A. K. (2005). The induction of the otic placode.
In M. W. Kelley, D. K. Wu, A. N. Popper, & R. R. Fay
(Eds.), Development of the ear (pp. 10–42). New York, NY:
Springer-Verlag.
Hamburger, V., & Hamilton, H. L. (1951). A series of normal
stages in the development of the chick embryo. Journal of
Morphology, 88, 49092.
Hard, E., & Larsson, K. (1975). Development of righting in
rats. Brain and Behavioral Evolution, 11, 53–59.
Herman, R., Maulucci, R., & Stuyck, J. (1982). Development
and plasticity of visual and vestibular generated eye
movements. Experimental Brain Research, 47, 69–78.
Heywood, P., Pujol, R., & Hilding, D. A. (1976). Development
of the labyrinthine receptors in the guinea pig, cat and
dog. Acta Oto-Laryngologica, 82, 359–367.
Hirabayashi, S., & Iwasaki, Y. (1995). Developmental perspec-
tive of sensory organization on postural control. Brain &
Development, 17, 111–113.
Horak, F. B., Schumway-Cook, A., Crowe, T. K., & Black, F. O.
(1988). Vestibular function and motor proficiency of children with impaired hearing, or with learning disability
and motor impairments. Developmental Medicine and Child
Neurology, 30, 64–79.
Hudspeth, A. J. (1982). Extracellular current flow and the site
of transduction by vertebrate hair cells. Journal of Neurosci-
ence, 2(1), 1–10.
Hudspeth, A. J. (1985). The cellular basis of hearing: The bio-
physics of hair cells. Science, 230, 745–752.
Hurley, K., M., Gaboyard, S., Zhong, M., Price, S. D., Wooltor-
ton, J. R. A., Lysakowski, A., . . . Eatock, R, K. (2006). M-like
K+ currents in type I hair cells and calyx afferent endings
38 BAlAnCE FunCtion AssEssmEnt And mAnAgEmEnt
https://t.me/medicina_free
of the developing rat utricle. Journal of Neuroscience, 26(40),
10253–10269.
Iwasaki, S., Chihara, Y., Komuta, Y., Ito, K., & Sahara, Y. (2008).
Low-voltage-activated potassium channels underlie the
regulation of intrinsic firing properties of rat vestibular
ganglion cells. Journal of Neurophysiology, 100, 2192–2204.
Jeffery, N., & Spoor, F. (2004). Prenatal growth and develop-
ment of the modern human labyrinth. Journal of Anatomy,
204, 71–92.
Jones, C., & Chen, P. (2007). Planar cell polarity signaling in
vertebrates. BioEssays, 29, 120–132.
Jones, S. M., Erway, L. C., Bergstrom, R. A., Schimenti, J. C., &
Jones, T. A. (1999). Vestibular responses to linear acceleration are absent in otoconia-deficient C57BL/6JEi-het mice.
Hearing Research, 135, 56–60.
Jones, S. M., & Jones, T. A. (2000a). Ontogeny of vestibular
compound action potentials in the domestic chicken. Journal
of the Association for Research in Otolaryngology, 1, 232–242.
Jones, T. A., & Jones, S. M. (2000b). Spontaneous activity in
the statoacoustic ganglion of the chicken embryo. Journal
of Neurophysiology, 83, 1452–1468.
Jones, S. M., & Jones, T. A. (2011). Genetics, embryology and
development of auditory and vestibular systems. San Diego,
CA: Plural Publishing.
Jones, T. A., Jones, S. M., & Hoffman, L. F. (2008). Resting
discharge patterns of macular primary afferents in otoconia-deficient mice. Journal of the Association for Research in
Otolaryngology, 9, 490–505.
Jones, C., Roper, V., Foucher, I., Qian, D., Banizs, B., Petit, C.,
. . . Chen, P. (2008). Ciliary proteins link basal body polarization to planar cell polarity regulation. Nature Genetics,
40(1), 69–77.
Kalluri, R., Xue, J., & Eatock, R. A. (2010). Ion channels set
spike timing regularity of mammalian vestibular afferent
neurons. Journal of Neurophysiology, 104, 2034–2051.
Kaufman, M. H. (1992). The atlas of mouse development. Amster-
dam, Netherlands: Elsevier Academic Press.
Kazmierczak, P., Sakaguchi, H., Tokita, J., Wilson-Kubalek, E.
M., Milligan, R. A., Müller, U., & Kachar, B. (2007). Cadherin 23 and protocadherin 15 interact to form tip link filaments in sensory hair cells. Nature, 449(7158), 87–91.
Kelley, M. W., & Chen, P. (2007). Shaping the mammalian
sensory organ by the planar cell polarity pathway. Inter-
national Journal of Developmental Biology, 51, 535–547.
Kenyon, G. S. (1988). Neuro-otological findings in normal
children. Journal of the Royal Society of Medicine, 81, 644–648.
Kros, C. J., Ruppersberg, J. P., & Rüsch, A. (1998). Expression
of a potassium current in inner hair cells during development of hearing in mice. Nature, 394, 281–284.
Ktrejcova, H. (1975). Proceedings: Vestibulometric evaluation
of the caloric reaction in children and in adults. Electroen-
cephalography and Clinical Neurophysiology, 39, 441.
Lanford, P. J., Lan, Y., Jiang, R., Lindsell, C., Weinmaster,
G., Gridley, T, . . . Kelley, M. W. (1999). Notch signaling
pathway mediates hair cell development in mammalian
cochlea. Nature Genetics, 21, 289–292.
Lanford, P. J., Shailam, R., Norton, C. R., Gridley, T., & Kelley,
M. W. (2000). Expression of Math1 and HES5 in the cochlea
of wildtype and Jag2 mutant mice. Journal of the Association
for Research in Otolaryngology, 1, 161–171.
Laouris, Y., Kalli-Laouri, J., & Schwartze, P. (1990). The post-
natal development of the air-righting reaction in albino
rats. Quantitative analysis of normal development and the
effect of preventing neck-torso and torso-pelvis rotations.
Behavioural Brain Research, 37, 37–44.
Lee, Y. S., Liu, F., & Segil, N. (2006). A morphogenetic wave of
p27kip1 transcription directs cell cycle exit during organ
of Corti development. Development, 133, 2817–2826.
Levens, S. L. (1988). Electronystagmography in normal chil-
dren. British Journal of Audiology, 22, 51–56.
Levi-Montalcini, R. (1987). The nerve growth factor: 35 years
later. Science, 237, 1154–1162.
Levi-Montalcini, R., & Angeletti, P. U. (1968). Nerve growth
factor. Physiological Reviews, 48(3), 534–569.
Lewin, G. R., & Barde, Y. A. (1996). Physiology of the neuro-
trophins. Annual Review of Neuroscience, 19, 289–317.
Li, G. Q., Meredith, F. L., & Rennie, K. J. (2010). Development
+
of K
and Na+ conductances in rodent postnatal semicircular canal type I hair cells. American Journal of Physiology
— Regulatory Integrative and Comparative Physiology, 298,
R351–R358.
Liebl, D. J., Tessarollo, L., Palko, M. E., & Parada, L. F. (1997).
Absence of sensory neurons before target innervation in
brain-derived neurotrophic factor-, neurotrophin 3-, and
Trk-C deficient embryonic mice. Journal of Neuroscience,
17(23), 9113–9121.
Lim, D. J. (1984). The development and structure of otoconia.
In I. Friedman & J. Ballantyne (Eds.), Ultrastructural atlas
of the inner ear (pp. 245–269). London, UK: Butterworth.
Lim, D. J., & Anniko, M. (1985). Developmental morphology
of the mouse inner ear. Acta Oto-Laryngologica, Supplement
422, 1–69.
Lim, D. J., & Rueda, J. (1992). Structural development of the
cochlea. In R. Romand (Ed.), Development of the auditory
and vestibular systems 2 (pp. 33–58). Amsterdam, Netherlands: Elsevier Science.
Lopez, C., & Blanke, O. (2011). The thalamocortical vestibular
system in animals and humans. Brain Research Reviews, 67,
119–146.
Ma, Q., Anderson, D. J., & Fritzsch, B. (2000). Neurogenin 1
null mutant ears develop fewer, morphologically normal
hair cells in smaller sensory epithelia devoid of innervation. Journal of the Association for Research in Otolaryngology,
1, 129–143.
Marcotti, W., Johnson, S. J., Holley, M. C., & Kros, C. J. (2003a).
Developmental changes in the expression of potassium
currents of embryonic, neonatal, and mature mouse inner
hair cells. Journal of Physiology, 548, 383–400.
Masetto, S., Bosica, M., Correia, M. J., Ottersen, O. P., Zucca,
G., Perin, P., . . . Valli, P. (2003). Na
+
currents in vestibular type I and type II hair cells of the embryo and adult
chicken. Journal of Neurophysiology, 90, 1266–1278.
Masetto, S., Perin, P., Malusà, A., Lucca, G., & Valli, P. (2000).
Membrane properties of chick semicircular canal hair cells
in situ during embryonic development. Journal of Neuro-
physiology, 83, 2740–2756.
2. ontogEny oF tHE vEstiBulAr systEm And BAlAnCE 39
https://t.me/medicina_free
Mbiene, J. P., Favre, D., & Sans, A. (1984). The pattern of
ciliary development in fetal mouse vestibular receptors.
Anatomy and Embryology, 170, 229–238.
Mbiene, J. P., Favre, D., & Sans, A. (1988). Early innervation
of hair cells in the vestibular epithelia of mouse embryos:
SEM and TEM study. Anatomy and Embryology, 177,
331–340.
Mbiene, J. P., & Sans, A. (1986). Differentiation and matura-
tion of the sensory hair bundles in the fetal and postnatal
vestibular receptors of the mouse: A scanning microscopy
study. Journal of Comparative Neurology, 254, 271–278.
McKenzie, E., Krupin, A., & Kelley, M. W. (2004). Cellular
growth and rearrangement during the development of
the mammalian organ of Corti. Developmental Dynamics,
229, 802–812.
Melagrana, A., D’Agostino, R., Pasquale, G., & Taborelli, G.
(1996). Study of labyrinthine function in children using
the caloric test: our results. International Journal of Pediatric
Otorhinolaryngology, 37, 1–8.
Montandon, P., Gacek, R. R., & Kimura, R. S. (1970). Crista
neglecta in the cat and human. Annals of Otology, Rhinol-
ogy, and Laryngology, 79, 105–112.
Montcouquiol, M., Rachel, R. A., Lanford, P. J., Copeland, N.
G., Jenkins, N. A., & Kelley, M. W. (2003). Identification of
Vang12 and Scrb1 as planar polarity genes in mammals.
Nature, 423, 173–177.
Montcouquiol, M., Sans, N., Huss, D., Kach, J., Dickman, J.
D., Forge, A., . . . Kelley, M. W. (2006). Asymmetric localization of Vang12 and Fz3 indicate novel mechanisms for
planar cell polarity in mammals. Journal of Neuroscience,
26(19), 5265–5275.
Morsli, H., Choo, D., Ryan, A., Johnson, R., & Wu, D. K.
(1998). Development of the mouse inner ear and origin of
its sensory organs. Journal of Neuroscience, 18(9), 3327–3335.
Morsli H., Tuorto, F., Choo, D., Postiglione, M. P., Sime-
one, A., & Wu, D. K. (1999). Otx1 and Otx2 activities are
required for the normal development of the mouse inner
ear. Development, 126, 2335–2343.
Mulch, G., & Petermann, W. (1979). Influence of age on
results of vestibular function tests. Review of literature
and presentation of caloric test results. Annals of Otology,
Rhinology, and Laryngology, 88, 1–17.
Nakahara, H., & Bevelander, G. (1979). An electron micro-
scope study of crystal calcium carbonate formation in the
mouse otolith. Anatomical Record, 193, 233–242.
Nayak, G. D., Ratnayaka, H. S. K., Goodyear, R. J., & Rich-
ardson, G. P. (2007). Development of the hair bundle and
mechanotransduction. International Journal of Developmen-
tal Biology, 51, 597–608.
Neef, J., Gehrt, A., Bulankina, A. V., Meyer, A. C., Riedel, D.,
Gregg, R. G., . . . Moser, T. (2009). The Ca
β2 regulates Ca
2+
channel abundance and function in inner
2+
channel subunit
hair cells and is required for hearing. Journal of Neurosci-
ence, 29(34), 10730–10740.
Nenov, A. P., Norris, C., & Bobbin, R. P. (1996). Acetylcholine
response in guinea pig outer hair cells. II. Activation of
a small conductance Ca
2+
-activated K+ channel. Hearing
Research, 101, 149–172.
Nichols, D. H., Pauley, S., Jahan, I., Beisel, K. W., Millen, K. J.,
& Fritzsch, B. (2008). Lmx1a is required for segregation of
sensory epithelia and normal ear histogenesis and morphogenesis. Cell and Tissue Research, 334(3), 339–358.
Nishida, Y., Rivolta, M. N., & Holley, M. C. (1998). Timed
markers for the differentiation of the cuticular plate and
stereocilia in hair cells from the mouse inner ear. Journal of
Comparative Neurology, 395, 18–28.
Ohyama, T., Groves, A. K., & Martin, K. (2007). The first
steps toward hearing: Mechanisms of otic placode induction. International Journal of Developmental Biology, 51,
463–472.
Ornitz, O. M., Atwell, C. W., Walter, D .O., Hartmann, E. E.,
& Kaplan, A. R. (1979). The maturation of vestibular nystagmus in infancy and childhood. Acta Oto-Laryngologica,
88, 244–256.
Parrad, P. J., & Cottereau, P. (1977). Apparition des réactions
rotatoires chez le rat nouveau-né. Physiology and Behavior,
18, 1017–1020.
Pauley, S., Matei, V., Beisel, K. W., & Fritzsch, B. (2005). Wir-
ing the ear to the brain: The molecular basis of neurosensory development, differentiation, and survival. In
M. W. Kelley, D. K. Wu, A. N. Popper, & R. R. Fay (Eds.),
Development of the inner ear (pp. 85–121). New York, NY:
Springer-Verlag.
Peterka, R. J., & Black, F. O. (1990). Age-related changes in
human posture control: Sensory organization tests. Journal
of Vestibular Research, 1, 73–85.
Peterka, R. J., Black, F. O., & Schoenhoff, M. B. (1990a).
Age-related changes in human vestibulo-ocular reflexes:
Sinusoidal rotation and caloric tests. Journal of Vestibular
Research, 1, 49–59.
Peterka, R. J., Black, F. O., & Schoenhoff, M. B. (1990b). Age-
related changes in human vestibulo-ocular and optokinetic reflexes: Pseudorandom rotation tests. Journal of
Vestibular Research, 1, 61–71.
Peterson, M. L., Christou, E., & Rosengren, K. S. (2006). Chil-
dren achieve adult-like sensory integration during stance
at 12-years-old. Gait & Posture, 23, 455–463.
Pickles, J. O., Comis, S. D., & Osborne, M. E. (1984). Cross
links between stereocilia in the guinea pig organ of Corti,
and their possible relation to sensory transduction. Hear-
ing Research, 15(2), 103–112.
Porter, J. D., & Balaban, C. D. (1997). Connections between
the vestibular nuclei and brain stem regions that mediate
autonomic function in the rat. Journal of Vestibular Research,
7, 63–76.
Pujol, R., Lavigne-Rebillard, M., & Lenoir, M. (1998). Devel-
opment of sensory and neural structures in the mammalian cochlea. In E. W. Rubel, A. N. Popper, & R. R. Fay
(Eds.), Development of the auditory system (pp. 146–192).
New York, NY: Springer-Verlag.
Rida, P. C. G., & Chen, P. (2009). Line up and listen: Planar cell
polarity regulation in the mammalian inner ear. Seminars
in Cell & Developmental Biology, 20, 978–985.
Romand, R., & Dauzat, M. (1982). Modification of spontane-
ous activity in primary vestibular neurons during development in the cat. Experimental Brain Research, 45, 2658.
40 BAlAnCE FunCtion AssEssmEnt And mAnAgEmEnt
https://t.me/medicina_free
Ruben, R. J. (1967). Development of the inner ear of the
mouse: A radioautographic study of terminal mitoses
[Supplemental material]. Acta Oto-Laryngologica, 220, 5–44.
Rüsch, A., & Eatock, R. A. (1996). A delayed rectifier conduc-
tance in type I hair cells of the mouse utricle. Journal of
Neurophysiology, 76, 995–1004.
Rüsch, A., Lysakowski, A., & Eatock, R. A. (1998). Postnatal
development of type I and type II hair cells in the mouse
utricle: Acquisition of voltage-gated conductances and
differentiated morphology. Journal of Neuroscience, 18,
7487–7501.
Salamat, M. S., Ross, M. D., & Peacor, D. R. (1980). Otoconia
formation in the fetal rat. Annals of Otology, Rhinology, and
Laryngology, 89, 229–238.
Sans, A., & Chat, M. (1982). Analysis of temporal and spatial
patterns of rat vestibular hair cell differentiation by tritiated thymidine radiography. Journal of Comparative Neurol-
ogy, 206, 1–8.
Sans, A., & Dechesne, C. (1985). Early development of vestib-
ular receptors in human embryos: An electron microscopic
study [Supplemental material]. Acta Oto-Laryngologica,
423, 51–58.
Schneider-Maunoury, S., & Pujades, C. (2007). Hindbrain sig-
nals in otic regionalization: Walk on the wild side. Interna-
tional Journal of Developmental Biology, 51, 495–506.
Schweizer, F. E., Savin, D., Suu, C., Sultemeier, E. R., &
Hoffman, L. F. (2009). Distribution of high-conductance
calcium-activated potassium channels in rat vestibular
epithelia. Journal of Comparative Neurology, 517, 134–145.
Shailam, R., Lanford, P. J., Dolinsky, C. M., Norton, C. R.,
Gridley, T., & Kelley, M. W. (1999). Expression of proneural and neurogenic genes in the embryonic mammalian
vestibular system. Journal of Neurocytology, 28, 809–819.
Shall, M. S. (2009). The importance of saccular function to
motor development in children with hearing impairments. International Journal of Otolaryngology, 10, 1–5.
Sher, A. E. (1971). The embryonic and postnatal development
of the inner ear of the mouse. Acta Oto-Laryngologica Sup-
plement, 285, 1–77.
Shiroyama, T., Kayahara, T., Yasui, Y., Nomura, J., & Nakano,
K. (1999). Projections of the vestibular nuclei to the thalamus in the rat: A Phaseolus vulgaris leucoagglutinin
study. The Journal of Comparative Neurology, 407, 318–332.
Silos-Santiago, I., Fagan, A. M., Garber, M., Fritzsch, B., &
Barbacid, M. (1997). Severe sensory deficits but normal
CNS development in newborn mice lacking TrkB and
TrkC tyrosine protein kinase receptors. European Journal of
Neuroscience, 9, 2045–2056.
Simmons, D. D. (2002). Development of the inner ear efferent
system across vertebrate species. Journal of Neurobiology,
53, 228–250.
Simmons, D., Duncan, J., Craponde Caprona, D., & Fritzsch,
B. (2011). Development of the inner ear efferent system. In
D. K. Ryugo, R. R. Fay, & A. N. Popper (Eds.), Auditory and
vestibular efferents (pp. 187–216). New York, NY: Springer.
Simon, D. B., Bindra, R. S., Mansfield, T. A., Nelson-Williams,
C., Mendonca, E., Stone, R., . . . Lifton, R. P. (1987). Muta-
tions in the chloride channel gene, CLCNKB, cause Bartter’s syndrome type III. Nature Genetics, 17, 171–178.
Sokolowski, B. H. M., Stahl, L. M., & Fuchs, P. A. (1993). Mor-
phological and physiological development of vestibular
hair cells in the organ-cultured otocyst of the chick. Devel-
opmental Biology, 155, 134–146.
Staller, S. J., Goin, D. W., & Hildebrandt, M. (1986). Pediatric
vestibular evaluation with harmonic acceleration. Otolar-
yngology–Head and Neck Surgery, 95, 471–476.
Sulik, K. K., & Cotanche, D. A. (2004). Embryology of the ear.
In H. V. Toriello, W. Reardon, & R. J. Gorlin (Eds.), Heredi-
tary hearing loss and its syndromes (2nd ed., pp. 17–36). New
York, NY: Oxford University Press.
Tibbling, L. (1969). The rotatory nystagmus response in chil-
dren. Acta Oto-Laryngologica, 68, 459–467.
Tierney, T. S., Russell, F. A., & Moore, D. R. (1997). Susceptibil-
ity of developing cochlear nucleus neurons to deafferentation-induced death abruptly ends just before the onset
of hearing. Journal of Comparative Neurology, 378, 295–306.
Valente, M. (2007). Maturational effects of the vestibular sys-
tem: A study of rotary chair, computerized dynamic posturography, and vestibular evoked myogenic potentials
with children. Journal of the American Academy of Audiol-
ogy, 18, 461–481.
Van Cleave, S., & Shall, M. S. (2006). A critical period for the
impact of vestibular sensation on ferret motor development. Journal of Vestibular Research, 16, 179–186.
Van der Laan, F. L., & Oosterveld, W. J. (1974). Age and ves-
tibular function. Aerospace Medicine, 45, 540–547.
Van de Water, T. R. (1976). Effects of removal of the stato-
acoustic ganglion complex upon the growing otosysts.
Annals of Otology, Rhinology, and Laryngology, 85, 2–31.
Van de Water, T. R. (1984). Developmental mechanisms of
mammalian inner ear formation. In C. Berlin (Ed.), Hear-
ing science (pp. 49–170). San Diego, CA: College-Hill Press.
Van de Water, T. R., Anniko, M. Nordemar, H., & Wersall,
J. (1977). Embryonic development of the sensory cells in
macula utriculae of mouse. I’Institut National de la Sante et
de la Recherche Medicale, 68, 25–35.
Veenhof, V. B. (1969). The development of statoconia in mice.
Amsterdam-London, UK: N.V. North-Holland Publishing.
Viener-Wacher, S. R., Toupet, F., & Narcy, P. (1996). Canal and
otolith vestibulo-ocular reflexes to vertical and off vertical axis rotations in children learning to walk. Acta Oto-
Laryngologica, 116, 657–665.
Wang, J., Mark, S., Zhang, X., Qian, D., Yoo, S. J., Radde-
Gallwitz, K., . . .Chen, P. (2005). Regulation of polarized
extension and planar cell polarity in the cochlea by the
vertebrate PCP pathway. Nature Genetics, 37(9), 980–985.
Wang, Y., Guo, N., & Nathans, J. (2006). The role of Frizzled3
and Frizzled6 in neural tube closure and in the planar
polarity of inner-ear sensory hair cells. Journal of Neurosci-
ence, 26(8), 2147–2156.
Wiener-Vacher, S. R., Toupet, F., & Narcy, P. (1996). Canal and
otolith vestibule-ocular reflexes to vertical and off vertical axis rotations in children learning to walk. Acta Oto-
Laryngologica, 116, 657–665.
2. ontogEny oF tHE vEstiBulAr systEm And BAlAnCE 41
https://t.me/medicina_free
Woods, C., Moncouquiol, M., & Kelley, M. W. (2004). Math1
regulates the development of the sensory epithelium in
the mammalian cochlea. Nature Neuroscience, 7, 1310–
1318.
Xiang, M., Gao, W. Q., Hasson, T., & Shin, J. J. (1998). Require-
ment for Brn-3c in maturation and survival, but not in fate
determination of inner ear hair cells. Development, 125,
3935–3946.
Xu, Z., Ricci, A. J., & Heller, S. (2009). Rethinking how hearing
happens. Neuron, 62, 305–307.
Yates, B. J. (1996). Vestibular influences on the autonomic ner-
vous system. Annals of New York Academy of Sciences, 781,
458–473.
Yates, B. J., & Miller, A. D. (1998). Physiological evidence that
the vestibular system participates in autonomic and respiratory control. Journal of Vestibular Research, 8, 17–25.
https://t.me/medicina_free
3
https://t.me/medicina_free
Practical Anatomy and Physiology
of the Ocular Motor System
Scott D. Z. Eggers
FUNCTIONAL CLASSES OF EYE MOVEMENTS
The common goal of all eye movements is to facilitate
a clear and stable view of the environment (Leigh &
Zee, 2006). Lateral-eyed animals like rabbits have
a large field of view to survey the environment and
avoid predators. But the brain cannot manage the data
processing requirements of a visual system with high
resolution across the entire visual field. Thus, lateraleyed animals generally have the tradeoff of poor visual
acuity. Birds of prey have evolved a visual compromise
by restricting their visual field in return for superior
visual acuity within that narrow field. Other frontaleyed animals like humans have developed a small area
of very high spatial resolution at the center of the retina
(the fovea) while maintaining lower resolution in the
periphery. This “foveal compromise” (Wong, 2008)
solves the problem of information overload but also
requires that the image of an object of interest fall on
the fovea for maximal visual acuity.
Mechanisms have evolved to complement this
foveal compromise strategy by ensuring that images of
interest are brought to and maintained on the foveae of
both eyes. Image stability on the retina must be maintained despite object or head motion, because image
“slip” across the retina or movement away from the
fovea leads to blur or degrades visual acuity. Thus, one
category of eye movements helps hold target images
steady on the retina. These include (1) active processes
of the visual fixation system to hold the image of sta-
tionary objects on the fovea when the head is still;
(2)
the vestibular system to hold target images steady
on the retina during brief head movements; and (3) the
optokinetic system to hold target images steady on the
retina during sustained head rotation. A second category of eye movements has evolved to direct the highresolution fovea to objects of interest. These include
(1) the saccadic system to bring a target image rapidly
onto the fovea; (2) the smooth pursuit system to hold
the image of a small moving target on the fovea; and
(3) the vergence system to move both eyes in opposite
directions (i.e., convergence or divergence) in order to
simultaneously place the target image onto both foveae
regardless of target distance or eccentricity (Table 3–1).
To fulfill these visual requirements, the ocular motor
system requires complex anatomy and physiology at
every level, from the extraocular muscles to cortical
ocular motor regions.
Visual Fixation
The fixation system holds the image of a stationary
target on the fovea while the head is still. Fixation
may be a special type of smooth pursuit (suppressing
unwanted drift of the eyes) or an independent fixation
system. Rather than simply the absence of visible eye
movements, visual fixation actually consists of constant
miniature movements not detectable to the naked eye,
with the illusion of steady fixation. Normal fixation
includes (1) microtremor (<0.01 degree, up to 150 Hz);
43
Соседние файлы в папке Библиотека им академика М.И. Перельмана