perienced consciously.
And as William James emphasized, concerned is not a search for liaison between brain as a
even a consciously learned response can become so
habituated that it submerges to the non-conscious level
and thereafter is taken care of quite automatically by the
nervous system, more or less on the same plane as genet-
ically endowed behavior.
Perception, memory, learning, and consciousness are
all biological activities of the nervous system. As Herrick
remarked: "The problem with which science is properly
Anatomical Basis of Attention
Mechanisms in Vertebrate Brains
M. E. SCHEIBEL a n d A. B. SCHEIBEL
A COMPLEX OF neurons and fibers that constitute the most
primitive part of the brain wanders through the substance
of the brain stem. This archaic system shows little evidence
of rigid structure, and approaches, in some respects, the
communication engineer's ideal of the statistical net. We
cannot call this reticular formation truly isotropic, but here
one can find no hint of the columnar arrays of cortical
pyramids, the file-on-file of bidimensional Purkinje cells
of cerebellum, nor even the whorled clusters of sensory
field cells, such as characterize analogically mapping fields
of sensory relays.
But perhaps as a result of this muted statement of its
structural theme, there resides within the brain-stem
core a remarkable pluripotentium of functional roles that
have become increasingly obvious over the past quarter
century and that probably mark it as the most critical
integrative center of the brain—determiner of operational
modes, gating mechanism for all sensory influx, modulator
and monitor of cortical function, readout mechanism for
the cortical differentiative and comparative processes and
gain manipulator for motor output. A brief,
chronologically oriented survey of the development of our
thoughts about this multi-faceted system may prove
useful in gaining insight into its present stature.
M. E. SCHEIBEL AND A. B. SCHEIBEL Departments of
Anatomy and Psychiatry and Brain Research Institute, UCLA,
Medical Center, Los Angeles, California
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
physical instrument and some other entity which is mind.
What we must do is to discover those characteristics of
brain as living tissue which enable it to have as one of its
own intrinsic physicalistic properties the awareness that
we call mind. . . . What we need here is more knowledge
of the actual structure and operation of the nervous mech-
anisms that do the thinking."55
Historical resume
"It is known from embryology that most of the leftover
cells of the brain stem and spinal cord which are not con-
cerned in the formation of motor root nuclei and purely
sensory relay nuclei are utilized in the production of the
formatio reticularis." In this curiously negative manner
commences a short and prescient article by W. F. Allen,'
published in 1932, which sketched out an envelope of
functional roles for the brain-stem reticular core that was
to be followed by investigators for the next three decades.
But the tenor of the sentence also reflects almost a century
of neurological prejudice, which conceived the reticular
core of the brain stem largely as filler—a kind of neuro-
ectodermally derived excelsior, in which were cushioned
the more functionally attractive cranial nerve nuclei.
During the past quarter century, there has been a dra-
matic change in value judgments. Compared to the spec-
trum of presently recognized reticular functions, ranging
from homeostat operations practiced on the internal mi-
lieu to modulation and control of highest psychic func-
tions, the workings of cranial nerve nuclei seem almost
humdrum. If our appreciation of the astonishing range of
physiological possibilities inherent in the core seems a
strictly contemporary phenomenon, it was not necessary
that it be so, for the anatomical and physiological litera-
ture has been replete with clues pointing in these directions
since the turn of the century.
Anatomical studies of Kohnstamm and Quensel,2,3
577
which suggested pooling of a number of afferent and ef-
ferent systems upon the reticular core, led them to propose
this area as a "centrum receptorium" or "sensorium commu
ne" - a common sensory pool for the neuraxis. Al-
though their attempts to interpret these data within a
clinical frame were not entirely convincing, implications of
their thesis included the integration of heterogeneous con-
vergent stimuli and an alternative centripetal path lashed
in parallel with the classical long sensory tracts—both
astonishingly contemporary conceptions.
At about the same time, Golgi chrome-silver studies by
Held4 and Cajal5 emphasized characteristics of input-out-
put systems and the possibilities for a spectrum of interac-
tions between reticular cells and intrinsic axonal systems.
Cajal called attention to the characteristic bifurcating axo-
nal output of a number of reticular neurons and their pro-
jection toward more rostral and caudal areas of the neu-
raxis. The caudal-flowing system was established as pro-
jecting upon spinal cord by retrograde studies of Kohn-
stam6 and van Gehuchten,7 and by anterograde (Marchi)
studies in the hands of Lewandowsky.8 However, the clas
FIGURE 1 Schematic representation of the relations of the
reticular core of the brain stem (black) with other systems of
the brain. Collaterals pour in from long sensory and motor
tracts (thin lines) and from the cerebral hemispheres (arrows
directed downward into reticular core). The core acts in turn
on cerebral and cerebellar cortices (upward-directed arrows)
578 BRAIN ORGANIZATION
sic Marchi study was that of Papez,9 who described three
major descending (rcticulo-spinal) tracts, their apparent
origins in the brain stem, and their approximate terminal
locations in the cord. This work, later to be amplified by
anatomical studies of Niemer and Magoun,10 Torvik and
Brodal,11 and Nyberg-Hansen,12 provides the hodological (
fiber bundle or tract) substrate for reticular modulation of
spinal mechanisms.
The studies of Allen" offered a highly intuitive preview
of the physiological role of the core in modulating rostral
and caudal structures. Allen was primarily an anatomist,
so his conclusions were largely speculative. Furthermore,
the time was not ripe for such ideas, and his paper was to
be forgotten for over a decade.
Equally pregnant was the discovery of Bremer,13,14
made only three years later, regarding the relation of brain
stem to cortex. He noticed that after decerebration the
electrocorticogram of the resultant "cerveau isolé" prepara-
tion swung into and maintained a high-voltage, slow-
wave rhythm that was maintained as long as the animal
survived. Such rhythms had just been identified by
and associated structures, on spinal cord (long, downward-
directed arrow), and on central sensory relays (striped ar-
rows). (From F. Worden, and R. Livingstone, 1961. Brain
stem reticular formation, in Electrical Stimulation of the
Brain [D. Sheer, editor], Univ. of Texas Press, Austin, pp.
263-276.)
Loomis, Harvey, and Hobart" and by Gibbs, Davis, and
Lennox" as consonant with sleep; so Bremer was able to
reach a conclusion of signal importance to experimental
neurology. The cerebral cortex needs tonic sensory influx
to maintain the wakeful state and, by implication, one of
the important roles of the long sensory tracts was to main-
tain, via continuous stimulus bombardment, a state of cor-
tical—and accordingly, organismic—vigilance. The bril-
liance of this interpretation was scarcely marred by
Bremer's error in identifying the agency involved.
The group around Alexander Forbes was the first to es-
tablish some of the physiological characteristics of the
rostral end of the extra-lemniscal (reticular) system. A sig-
nificant series of papers spanning the period 19361943 de-
- interface in the rostral pons separating areas apparently
limited its probable location within the thalamus, the
widespread nature of its projection upon cortex, and the
characteristic pattern of recruitment waves that followed
its stimulation at low frequency.17'18 Following the war,
these studies were expanded by Jasper and his associates."-"
who defined in greater detail the probable anatomical
paths and electrophysiological characteristics of this sys-
tem. An interpretative capstone was placed on this grow-
ing body of data with the suggestion by Penfield22 that the
diencephalic intralaminar system might well serve as a " stations by reticular collaterals. Notwithstanding a recent
centrencephalon" for all neural activity. The classical
teachings ofHughlings Jackson23 as to the ultimate status of
the cerebral cortex in the neuraxial hierarchy was thus
supplanted. It had now become penultimate to the upper
end of the reticular core, and the principle of thalamo-
cortico-thalamic circulation already proposed by Cajal,'
Campion," and by Dusser de Barenne and McCulloch25
gained support and experimental verification.
In the meantime, the range of autonomic control exert-
ed by brain-stem reticular mechanisms had been investi-
gated by Ranson and his colleagues,26.27 thereby extending
the work of early investigators in this area, and culminat-
ing in concepts of overlapping medullo-pontine fields sub-
serving respiratory and circulatory patterns. Out of this
developed the observations of Magoun and Rhines28-3° on
brain-stem reticular override (inhibition and facilitation)
of ongoing spinal motor activity, with subsequent con-
ceptual modifications by Sprague and Chambers." Short-
ly thereafter followed the epochal report of Moruzzi and
Magoun,32 describing reticular control over cortical ac-
tivity and its relation to the spectrum of conscious states
from attentive awareness to deep sleep. These workers at-
tributed such effects to ascending polysynaptic conduction
over successive chains of short-axoned cells. That these
phenomena, in reality, depended importantly on high-
priority conduction in oligosynaptic and, in some cases,
monosynaptic channels to cortex was suggested by our
Golgi studies33 and more recently by intracellular analyses
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
of Magni and Willis."
Results of selective destruction of lemniscal and extra -
lemniscal (reticular) systems by Magoun and a group of
collaborators35,36 now placed the original findings ofBremer","
in correct perspective, as it became increasingly clear that
extralemniscal, rather than direct sensory-tract volleys,
were crucial to maintenance of cortical tonus. By
implication, sleep remained a passive process consequent
to decrementing levels of reticular activity, despite earlier
studies by Hess,37,33 which had shown that it could be ob-
tained by stimulation of appropriate brain-stem sites. It re-
mained for Jouvet33," and Rossi41,42 to document a complex
of interrelated states and to focus attention on an anatomical
productive of rhombencephalic (activated) sleep from
classical slow-wave sleep (see Jouvet, this volume).
Still another significant facet of reticular-core activities
has been revealed by Galambos43,44 and Hernandez-Pe6n,45, 48
who have been able to relate the size of sensory evoked
potentials to the focus of active interest momentarily ex-
pressed by implanted, freely mobile animal. Our own
Golgi investigations33,47 have revealed a suitable substrate in
the penetration of most first- and second-order sensory
disclaimer by Worden and Marsh,'" it is generally con-
ceded that, aside from modulation of tonic and phasic
components that comprise the activated state,'" the reticu-
lar core mediates specific delimitation of the focus of con-
sciousness with concordant suppression of those sensory
inputs that have been temporarily relegated to a secondary
role.
Knowledge of the intimate physiology of the core has
been gleaned from extracellular microelectrode studies by
a number of investigators,'°-52 and most recently by a
limited number of intracellular analyses34,53," that have
examined in some detail the problems of convergence of
sensory signals, habituation (response attenuation) of in-
dividual units to iterative stimuli,55.56 and a cyclical alter-
nation of unit sensitivity to the stimulus array.57 At the
same time, a group of Golgi studies has revealed much
about the relevant substrate,33,58-" and in providing circuit
paradigms for a system with this order of functional com-
plexity, has also suggested program modes applicable to
computer design.62
The core as a mosaic
In the tradition of the great cortical cytoarchitectonicists (
i.e., Brodmann," the Vogts," von Economo," etc.), a
number of attempts have been made to subdivide the re-
ticular core of the brain stem into component nuclei on
579
the basis of its appearance after aniline staining. The most
detailed essay in this direction is that of Olszewski and his
collaborators," who have identified more than 40 nu-
clear entities in man, and almost as many in rabbit, in the
interval between the spino-medullary junction and the
rostral mesencephalon. The most obvious advantage of
such a system is the moderate degree of uniformity pro-
vided in an invariant vocabulary for descriptions of vari-
ous portions of the core. Thus, all who are acquainted
with Olszewski's atlas will visualize approximately the
same sector when asked to consider, for example, the nu-
cleus reticularis pontis caudalis. However, the original
bases for cytoarchitectonic identifications are often ar-
bitrary, and individual judgments vary as to where a pat-
tern of somal morphology or distribution becomes suf-
ficiently different to merit recognition as another nucleus.
The problem is particularly vexing in the reticular core,
where obvious transitions are the exception rather than
the rule, and cells of many sizes are intermingled.
The significance of somal size and distribution can be
questioned as adequate criteria for isolation of a nuclear
field. In a complex territory like the brain stem, it is at best
an assumption that these factors will mirror differences in
input arrangements, efferent destinations, local specializa-
tion of the microchemical environment, presence of spe-
cific synaptic mediators, etc. A more likely disclaimer to
the validity of such parceling is the appearance of these
areas when impregnated by one of the Golgi modifica-
tions. The long, relatively unramified dendrites character-
istic of cells of the reticular core stream in all directions, es-
pecially when studied in transverse section. Such processes
transcend the cytoarchitectonically determined nuclear
limits, projecting into neighboring and even remote nu-
clear subdivisions. Examination of a number of species in-
dicates that the total extent of the dendritic domain of a
single cell may exceed one-third of the total cross-sectional
area of the stem.68 The processes of such a cell can easily in-
vade as many as half a dozen extraneous nuclear pools.
We may conclude that the apparent isolation of each
core nucleus from its neighbor is entirely an artifact or, al-
ternatively, that the relative segregation of cell bodies and
the mingling of dendrite shafts express fundamental dif-
ferences in the physiology of these entities (Purpura, this
volume). As is usual in cases such as this, truth will prob-
ably be found to lie somewhere between.
Dendritic apparatus
The morphology and patterning of dendrites has been a
subject of interest since the initial studies of Golgi.69 With
the exception of the lateral reticular nucleus (noyau du
cordon lateral'),dendrites of reticular core cells appear rela-
580 BRAIN ORGAN I ZAT ION
tively straight, long, and unramified. The uniqueness of
this pattern has been remarked by a number of investiga-
tors,5,70,71 and referred to as isodendritic by one group,"
who see in its similarity to archaic patterns a relation to in-
tegrative rather than differcntiative or comparative func-
tion. Such dendrites are characteristically thick (3 to 8 mi-
crons) at the somal junction, and may taper over a course
of several hundred microns or more to terminal segments
only a fraction of a micron in diameter. Golgi sections sug-
gest, and electron micrographs show, that the dendrite
tips may be quite as densely covered with presynaptic
structures as more proximal portions of the shafts. The
functional significance of terminal structures committed to
sections of postsynaptic membrane so distant from the
presumed spike-trigger area around the axon hillock re-
mains a question of more than theoretical interest (Pur-
pura, this volume, and Note 78).
The available dendritic (postsynaptic) membrane is
large, often constituting 85 to 95 per cent of the total re-
ceptive surface of the neuron." Such a figure agrees with
estimates of Sholl74 and others for the fraction of post-
synaptic membrane represented by the dendritic segment
of cortical neurons. The surface of these processes is irreg-
ular, often nodular, and unevenly covered by hairy pro-
tuberances, or "spines." The reality of the spiny apparatus
has been attested by electron micrographs by Gray" and
others, with the suggestion of elaborate intraspinous struc-
tures—at least in cortical spines—and a special population
of presynaptic junctions effected at the apex or base of the
spine. Although these structures are not so numerous (0.1
to 0.3 per linear micron) as in cortex (0.4 to 0.7 per linear
micron)," their presence is virtually invariant and indi-
cates an as-yet-unknown specialization of postsynaptic
membrane.
If the dendrites radiate widely, as seen in cross section,
their appearance is very different in sagittal planes. The
majority of reticular neurons over the medial two-thirds of
the core continue to show impressive degrees of spread in
the transverse dimension (dorso-ventral and latero-lateral),
but little or no projection in the rostro-caudal dimension.
The appearance is one of marked compression along the
long axis of the stem, resulting in a series of flattened den-
dritic domains, piled one on another like a stack of poker
chips. We originally suggested33 that this configuration in-
troduced a significant dimension of localization of input,
because each cell thereby "looked" only at a limited series
of inputs from a single level along the input continuum.
This seemed especially likely, as presynaptic components
invariably parallel the postsynaptic (dendritic) surface, and
virtually all terminating axonal elements pour into the
core at right angles to the long axis.
The modular nature of this "stack of chips" organiza-
 rostral
C. s. RI.
act:-

NV

C.

Tr.sP. 8

T.

N.mes.V-
10
p.Br N.r.I-

N.e e-

Tros
VP

N.Ir.sp.V
N.XII caudal

FIGURE 2 A series of equally spaced drawings of transverse
Nissl-stained sections through the brain stem of the cat to
show the grouping of cells in the reticular core. Various-
sized dots on the right indicate specific cell bodies, while
dotted lines on the left indicate approximate boundaries of
major reticular nuclei. Terminology and nuclear areas are
based on the studies of Olszewski in the rabbit, with minor
modifications. The following list of abbreviations refer
only to structures making up the reticular core: a,
Accessory group of paramedian reticular nucleus: d, Dorsal
group of paramedian reticular nucleus; h, region poor in
cells of Meesen and Olszewski surrounding motor
trigeminal nucleus; k, cell group k of Mecsen and
Olszewski; in, cell
group m of Meesen and Olszewski; N. ic., Nucleus inter-
calatus ; N. in., Nucleus interpositus; N. r. 1., Lateral reticu-
lar nucleus; N. r. p., Nucleus reticularis paramedianus ;
N. r. t., Nucleus reticularis tegmenti pontis; N. t. d., Dorsal
tegmental nucleus; N. t. v., Ventral tegmental nucleus;
P. g. Periaqueductal gray; P. h., Nucleus prepositus hypo-
glossi ; R. gc., Nucleus reticularis giganto-cellularis ;
Nucleus reticularis lateralis (of Meesen and Olszewski); R .
mes, Reticular formation of mesencephalon; R.n., Nucleus
of the raphe; R.p.c., Nucleus reticularis parvocellularis ; R.
p.o., Nucleus reticularis pontis oralis ; R.v., Nucleus
reticularis ventralis; v, Ventral group of paramedian reticu-
lar nucleus. (From Brodal, Note 80)

NEURAL SUBSTRATES OF ATTENTION MECHANISMS 581

 FIGURE 3 Cross section through upper medulla ofnew-born
kitten, stained by the Golgi method, showing the general
appearance of reticular cell bodies and dendrites at this level.
There is marked variation in size and shape of both den-
drites and cell bodies. The dendrite systems tend to radiate
in all directions in this plane of section and sometimes to in-
tional pattern has recently been exploited by Kilmer and
McCulloch,62 who have used it as a paradigm for a new
generation of computers, which will show increased de-
grees of flexibility in dealing with data influx, will habitu-
ate over time to iterative stimuli, etc. Although the con-
cept of a reticular core made up of a subinfinite number of
modules as numerous as its cellular complement may seem
overly divisive, it shows a rather good conceptual fit with
ideas emerging from single-unit analysis of the structure 5°_52
As we will stress again, most neurons appear to receive
upon their soma-dendrite surfaces an appreciable, but not
unlimited, convergent sampling of the many inputs af-
ferent to the core. Each neuron receives its own idiosyn-
cratic mix—in terms of afferent selection and loading—
from the total influx, so it has seemed appropriate to some
of us52 to consider each neuron as an integrating subcenter,
operating upon its own peculiar combination of afferents.
The sum of these neuronal subcenters would, in turn,
make up the total mosaic of the operating reticular core,
in which each element performed operations upon one
segment of the total envelope.
If we accept each reticular element as one of a family
582 BRAIN ORGANIZATION
vade a fiber bundle from which they receive excitation. Ab-
breviations : C. Rest., restiform body; n VIII (Deft.), Deiters
nucleus of the vestibular complex; n.VIII (Schw.),
Schwalbe's nulceus of the vestibular complex; MLF, medial
longitudinal fasciculus ; n.V.d., descending nucleus of the
trigeminal nerve; Tr. Pyr., pyramidal tract.
that samples a limited and idiosyncratic fraction of the
total reticular afferent supply, we might pause to con-
sider the relative significance of neuronal soma and den-
drite, respectively, in servicing the presynaptic array. The
widely ranging dendritic apparatus has already been seen
to penetrate areas rather remote from that of the parent cell
body, in quest of presynaptic excitation. We have suggested
elsewhere that the dendrites appear to point toward, and
reach for, potent sources of influx, which suggests that
trophic or neurobiotactic forces are operant during the de-
velopmental phase. We have previously shown evidence
of clear-cut segregation of afferent terminals on various
portions of dendrite shafts in cortex" and on spinal inter-
neurons." Although we do not have evidence of concen-
trated afferent terminal populations on various portions of
single reticular dendrites, there seems little doubt that
many reticular cell dendrites will show marked predomi-
nance of inputs, depending on the location of that shaft
relative to the spectrum of afferent sources (Figure 5).
Thus, for a cell located in the ventral-lateral quadrant of
medulla, it might be appropriate to refer to shafts that are
predominantly cortico-spinal loaded, spin-thalamic load-
 FIGURE 4 Sagittal section through the lower half of the
brain stem of a 10-day-old rat. Most of the dendrite mass of
reticular core cells is organized along the dorso-ventral axis
as seen in this type of section, with marked compression
along the rostro-caudal axis. This orientation places the
dendrites parallel to the terminal presynaptic components
which in this case arise from pyramidal tract (Tr. Pyr.) and
from a single axon of a magnocellular reticular neuron (n.
ed, descending trigeminal loaded, etc., thereby emphasiz-
ing the predominant population of afferents to be found
on that process.
However, this concept suffers some dilution because
each reticular cell lies in a matrix of fibers, largely (al-
though not entirely) core derived and projecting for vary-
ing distances in the rostro-caudal (and occasionally trans-
verse) direction. It is a characteristic property of such
axons to give off collaterals at right angles more or less
continuously along their path.33,60 These collaterals make
up a spectrum of lengths; some constitute little more than
minimal specializations of the fiber in transit (boutons-en-
passage). Each reticular dendrite must thread among the
paths of tens of thousands of such axons, so a considerable
number of axodendritic contacts must be expected from
this source, even if mere contiguity of nonspecialized axon
conductors with dendritic membrane is not in itself suffi-
cient for information transfer. It seems logical to expand
the hypothesis of sensory-annotated dendritic monads to
include in each case a sprinkling of core-derived afferents.
The moment-to-moment dendritic membrane loading
would thereby represent an integrate of specific and non-
specific inputs. In turn, the algebraic summation of such
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
retic. mag.). This type of dendrite organization, which is
especially characteristic of reticular cells of the medial 2/3 of
the core, produces sets of essentially two-dimensional mod-
ular neuropil fields leading to the stack of chips analogy (see
text and inset diagram at lower left). This is contrasted with
dendritic patterns in the adjacent hypoglossal nucleus (n.
XII); n. inf. ol., inferior olive; n. pontis, the pons. (From
Scheibel and Scheibel, Note 33)
states for each major dendrite shaft of a specific reticular-
core neuron would represent (with additions from the
somal synaptic scale) the level of synaptic drive being ap-
plied to the soma—initial segment at a specific point in
time.
It seems beyond question that the output of each reticu-
lar element represents a vector of this type; it therefore
follows that specific informational content, intrinsic to each of
the afferent sources, is lost in the integrative process. The out-
put of each unit must represent intensity only. It may not
be out of place to suggest the analogy that a major func-
tion of the core is to describe not the pageantry and color
of the passing parade, but the loudness of the shouting that
accompanies it.
The presypnatic influx
The reticular core of the brain stem may be thought of as
sitting athwart all incoming and outgoing information-
carrying systems. The collaterals and terminals that pour
in at every level receive a continuous stream of samples of
the activity ongoing in these tracts. In general, archaic af-
ferent and efferent systems, like the spino-thalamics and
583
 FIGURE 5 Semi-schematic cross section through upper third
of medulla, showing relations of a few presynaptic com-
ponents to dendrites and somata of reticular neurons. Some
dendrites appear to point toward the source of their most
significant presynaptic supply, individual fibers of which
may effect multiple terminations along a single dendrite
shaft. The nature of synaptic loading along each of a group
of dendrites radiating from a single soma may be unique.
Al-
extrapyramidals, are more thoroughly represented than
the phylogenetically newer tracts, such as the dorsal col-
umn—medial lemniscus and pyramid. The former (medial
lemniscus), in fact, probably contributes no collaterals to
the core, thus suggesting that information with a high de-
gree of locus and mode specificity is not crucial to the op-
eration of the reticular mosaic.
A fairly massive spino-reticular system introduces af-
ferent activity into medullo-pontine levels, along the long
axis of the tract, as do descending components in the cen-
tral tegmental fasciculus and the brachia efferent from
cerebellum. The most consistent afferent source, however,
appears to be the collaterals that pour in from sagittally
coursing fillets along the ventral and lateral aspects of the
stem and on each side of midline.5,33 In general, the ag-
gregate of reticulopetal fibers (fibers afferent to the reticu-
lar core) may be divided into several categories, which in-
584 BRAIN ORGANIZATION
though the apparent segregation of presynaptic supply is
very obvious in this sketch, the density of presynaptic ter-
minals in the core actually results in appreciable degrees of
convergence of heterogenous afferents upon each shaft.
Vestib. compl., vestibular nuclear complex; V. Desc., de-
scending trigeminal; Lem. sp., lemniscus spinalis (spino-
thalamic tract); Tr. Pyr., pyramidal tract; MLF, medial
longitudinal fasciculus.
dude: (1) fibers from more rostrally placed centers (cere-
bral cortex, basal ganglia, diencephalon, and limbic or al-
locortex) ; (2) fibers ascending from the spinal cord; (3) fi-
bers originating in the cerebellum and other brain-stem
structures such as the geniculate bodies, colliculi, etc.
These data are discussed in comprehensive form in several
places.79,80 Accordingly, only a few summarizing state-
ments will be made here.
THE CORTICO-RETICULAR PROJECTION This makes up
the most dramatic, if not the largest, descending system.
The fibers originate in an area centering on the sensor-
motor strip, but extend rather widely into adjacent corti-
cal areas, and terminate in "nucleus reticularis giganto-cel-
lularis, more rostally than caudally, while the pontine ter-
minal region is found in the nucleus reticularis pontis oralis
and the nucleus reticularis pontis caudalis, chiefly in its
rostral part."" Some terminations are also found in nu-
cleus reticularis tegmenti pontis.
Of particular interest are the components descending in-
directly from the limbic system upon midline tegmental
structures, a projection studied in some detail by Nauta.82
It seems clear that several routes are taken through thala-
mus and hypothalamus, with the eventual focus upon the
mesencephalic nuclear fields of Bechterew and Gudden,
from which arise an ascending system to complete a mas-
sive limbic-midline circuit. Informational by-products of
activity in this circuit may also be disseminated laterally
throughout the tegmentum and central tectum by the
somewhat enigmatic radiation first described by Weis-
schedel.83 We are as yet unable to decide from our material
whether this represents the fine-fibered lateral projection
of large numbers of small neurons situated in the peri-
aqueductal gray, or, in whole or in part, a projection of
neuroglial stalks, running from periependymal neuroglia
that line the Aqueduct of Sylvius. If the latter possibility
should be proved, it might introduce interesting possibili-
ties for neurohumoral interactions at this level of the core.
DESCENDING TECTAL COMPONENTS Originating in the
colliculi and pretectal area, these components terminate in
the same general areas as do cortico-reticular components,
probably as part of the tecto-bulbo-spinal correlation sys-
tem. The terminal patterns of reticulo-petal components
from basal ganglionar and diencephalic sites must still be
worked out completely, although Golgi material indicates
a hierarchy of fiber lengths and terminal stations through-
out the length of the core.'"
SPINO-RETICULAR FIBERS These fibers originate from all
levels of the cord,80,84 ascend in the ventro-lateral funiculus of
the cord and terminate along an extensive core of medullo-
pontine tissue, roughly congruent with areas of termi-
nation of the cortico-reticular projection. In addition, an
appreciable fraction of spino-reticular elements project
through, or terminate in, the lateral reticular nucleus,
which also receives many of the lower brain-stem termi-
nals of the spino-bulbo-thalamic system. This nucleus
projects, in turn, upon cerebellum, while the majority of
more centrally placed reticular fields (excluding para-
median nucleus and nucleus reticularis tegmenti pontis) re-
late back only to upstream or downstream centers."
THE CEREBELLO-RETICULAR CONTINGENT The con-
tingent enters the stem directly via the superior and in-
ferior cerebellar peduncles. (The former is found primarily
in the hooked bundle of Russell.) As summarized by Bro-
dal," major terminal stations in the reticular formation in-
clude nucleus reticularis tegmenti pontis and nucleus re-
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
ticularis pontis caudalis, the paramedian nucleus, and por-
tions of giganto-cellularis.
AFFERENT PROJECTION UPON THE RETICULAR CORE De-
tails can be studied in Golgi sections through the upper
medulla, which will serve as a paradigm. Figure 6 shows
that collaterals from long ascending and descending tracts (
i.e., spino-thalamic, descending trigeminal, vestibulo-
spinal, tecto-bulbo-spinal, cortico-spinal, etc.) turn into
the plane of section and terminate in overlapping patterns
within the reticular core. At a higher level of resolution (
Figure 7, right side), these terminals can be seen to cover
the soma and dendrites of most neurons within the field,
thereby making up the total roster of afferent supply or
synaptic scale85 upon the neuron.
The convergence of such a heterogeneous afferent array
can also be demonstrated physiologically by means of oc-
clusion techniques with macroelectrodes and by single-
unit recording methods with the extracellular or intracel-
lular microelectrode. The left side of Figure 7 shows strips
from a series of records illustrating the response of a single
unit to a group of disparate inputs, as well as its lack of re-
sponse to others. Data of this sort underline the extensive
but not unlimited convergence of inputs upon each reticu-
lar element. The unique quality of the synaptic "mix"
upon each neuron emphasizes the mosaic-like nature of the
core, with each neuron acting as a tiny integrating subcen-
ter within the complex.
Cyclic phenomena
Anatomical analysis of an input array spread over the
soma-dendrite complex of the average reticular neuron is
sharply and unequivocally stated when subjected to suc-
cessful Golgi staining or electron micrography. Further-
more, mechanical techniques that attempt the separation
of synaptic boutons from the subsynaptic membrane bear
witness to the tenacity with which presynaptic endings
cling to the postsynaptic ensemble. Such procedures, in
the hands of de Robertis,86,87 who has differentially cen-
trifuged the various fractions of crude separata down
through a series of sucrose interfaces, reveal numerous ele-
ments in the bouton fraction still clinging to fragments of
postsynaptic membrane. Clearly, this is no relationship
that can be reversibly severed by the swing of an astrocyte
tail, as was once hypothesized by Cajal.5
Yet, electrophysiological data that we have gathered
from long-term, extracellular recording from a number of
medullo-pontine reticular elements suggest that no matter
how anatomically stable the synaptic relationships appear
to be, their functional interactions are dynamic and cy-
clic.57 In about 80 per cent of the reticular elements from
585

FIGURE 6 Transverse section through the upper third of
the medulla ofa 10-day-old kitten, showing the convergence
and overlapping of terminating afferent fibers in the reticu-
lar core. On the left, a small group of fibers are drawn di-
rectly from the microscope; on the right, as a group of
overlapping sectors. Reading clockwise from the top, the
afferents include the descending periventricular system (Tr.
which we were able to record for periods of from 10 to 12
hours each, there was a succession of reactive phases dur-
ing which the cells appeared alternately sensitive to sen-
sory bombardment from the external milieu (mild sciatic
shocks of 1 to 2 volts of M millisecond, administered at 1-
per-second frequency) and from the internal milieu, as
exemplified by rhythms following the respiratory cycle.
During these respiration-sensitive periods, the reticular
elements were completely refractory to exteroceptive
stimuli (Figure 8). Similarly, during sciatic shock sensitive
phases, no traces of interoceptive-modeled activity were
ever seen. Each cell seemed to follow a unique temporal
pattern within a general order of magnitude of M to 3
hours (Figure 9). There was no discernible relation of these
swings to sleep-wakefulness cycles nor were there indica-
tions of general physiological changes serving as substrate
to the alterations.
Speculation on the mechanisms underlying this phe-
nomenon include hypothecated endocellular rhythms,
which periodically change the receptive sensitivities of the
586 BRAIN ORGANIZATION
f. Unc.
Tr. Periventr. Vestib. ,
desc. retic. //
/ / ,
Ii \
\ /
\ / 1./ /
1
Tr. Pyr.
Periventr. desc.); vestibulo-reticular fibers (Vestib. retic.) ;
the uncinate fasciculus from cerebellum (f. Unc.) ; the tract
of the descending fifth nerve (Tr. V desc.); lemniscus spinalis (
spinothalamic tract) (lernn. spin.); pyramidal tract (Tr.
Pyr.); and midline white matter, including medial reticulo- 3,4
spinal fibers (T. Rs. med., etc.). (From Scheibel and Scheibel,
Note 33)
subsynaptic mosaic (see notes 88 and 89 for empirical data
of possible relevance to this problem). An alternative sug-
gestion regarding mechanisms is the presence of a popula-
tion of pacemaker neurons that controls postsynaptic re-
sponse either through presynaptic manipulation90,91 or via
interaction of satellite oligoglia, which we have shown re-
ceive terminating collaterals from the same preterminals
that innervate the neuron.92 Each suggestion carries with it
its own intrinsic problems, the most intriguing of which
endow our speculations on the pacemaker.
In a cellular matrix as complex as the reticular forma--'
tion, if one hypothesizes a pacemaker elite that comprises a;,
fraction of the mass of followers, the quest for order sug-
gests a hyper-elite to pace the pacemakers, and so on to the
ultimate absurdity of one supreme, pontifical neuron. At
more attractive variant of this hypothesis involves mobile a
nd redundant command, which temporarily vests pacin
activities on those local arrays whose information loadin
is of a more biologically urgent nature than that of ado
jacent, more indecisive domains. As this content-mosai

varies through each fraction of time, it might be resona
ble to expect that centers of pacemaking activity would shift
widely through the core—a suggestion not out of context
with our own data already mentioned, which indicates
multi-modal operations of reticular elements.
If we assume that the relatively small fraction of reticul
ar elements from which we have led potentials is repres
entative of the entire array, and that periodically the ma-
jority of brain-stem-core neurons lower their threshold to
inputs, first of one sort and then of another, the significance
of such behavior becomes a matter of some concern. We
can only speculate that the cyclic activity which seems to
couple and uncouple neural elements from one circuit or
another may help preserve functional isotropicity within
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
tion; M and N : driven after a brief latency by single shocks
to cortex (note expanded time base beneath these last two
records). Strips C, D, E, and F show another unit that is sen-
sitive to nose pressure and can also be driven by clicks.
Cells A, B, and C are bulbar reticular elements in 10-day-
old kitten whose synaptic scale of terminating afferents is
partially shown. Horizontal-running fibers such as A, 1
through 7, and B, 6 through 8, may belong to long spino-
reticular and reticulo-reticular components while B, 1
through 4, represent sensory collaterals and cerebelloreticu-
lar terminals. (From Scheibel and Scheibel, Note 33)
the reticular net. It may well be that without periodic in-
put gating of this sort, masses of reticular elements would
gradually be drawn into the full-time service of that sys-
tem, sensory or motor, within whose geographic domain
they most nearly fall. It is conceivable that by such a proc-
ess of gradual acquisition and entrapment, increasing num-
bers of reticular elements would be lost to that range of
multi-potent functions that characterize core operations. A
neural ensemble, previously free to engage in multi-modal
integrative operations, could then conceivably become a
crazy quilt of satraps, each one only an extension of the
specific system demanding its fealty. Make break swings
-
of the sort we have described might effectively counter
such a trend.
587

FIGURE 8 Bulbar reticular unit showing responsiveness first to
endogenous, then to exogenous stimuli. Strips A through G
show a bulbar unit driven by respiratory activity but totally non-
responsive to sciatic stimulation (1/sec, 2 volts, 1/2 msec). Strip
H shows activity of the same unit 1/2 hour later. It is now highly
responsive to the same type of sciatic stimulation but shows no
evidence of being driven by respiratory activity. (From Scheibel
and Scheibel, Note 57)
588 BRAIN ORGANIZATION
The axonal outflow
The axonal outflow of the reticular core consists of a spec-
trum of conducting systems projecting in three main di-
rections—caudally upon spinal cord, rostrally upon sub-
cortical and cortical centers, and dorsally to cerebellum.
In addition, a vast and complex pattern is generated by the
output of the core operating upon itself through collat-
erals en passage. Axonal collaterals of greater length also
penetrate sensory and motor nuclei of cranial nerves en
route. The first three projections can be demonstrated with
some degree of rigor by the usual hodological methods
employing retrograde or anterograde degeneration. Using
modifications of the latter, Broda80 has shown that a system
of reticulo-cerebellar projections originates from the lateral
and paramedian nuclei and from the nucleus reticularis
tegmenti pontis, which differ appreciably in their terminal
stations. Use of the same technique has shown that the
reticulo-spinal projection arises from cells spread along the
medial two-thirds of medulla and pons, with particular
reference to nuclei pontis oralis et caudalis, nucleus
reticularis giganto-cellularis, and reticularis ventralis.
Similarly, ascending reticular axons identified by lesions
at the meso-diencephalic junction (Figure 10) arise from
the same general nuclear fields, suggesting either an in-
sa
it
 -0•
• R5 t.

Nt.d.
.

—F.I.m.
N.11

N.r.t.

j
i,

01
N.f.c.
N.c.e.
T.s.

N.m.X
Tr.spV
N.X1I
01.1

F I G U R E 1 0 A lesion at the meso-diencephalic junction of
cat brain stem and the position of cell bodies originally send-
ing rostral-coursing axons through the site as determined by
retrograde techniques (see text). Notice that a predominantly
unilateral lesion results in degenerating cell bodies (dots) on
both sides of the stem more caudally, and as far posteriorly
as the middle third of the medulla. S.n., substantia nigra ;
III, third nerve nucleus and nerve; Elm., medial longitudi-
nal fasciculus; N.r., nucleus ruber ; P, pons; C. s., superior
colliculus; IV, fourth nerve nucleus; N.r.t., nucleus re-
ticularis tegmenti; C.i., inferior colliculus; N. mes. V,
mesencephalic root of the fifth nerve; N.t.d., dorsal teg-
mental nucleus; N.1.1., nucleus of lateral lemniscus ; VI, sixth
nerve nucleus ; N VII, seventh nerve; N.n.V, sensory
trigeminal nucleus; N.c., cochlear nucleus; 01.s., superior
olive; P.h., perihypoglossal nucleus; N.f.c., cuneate nu-
cleus; N.c.e., external cuneate nucleus; T.s., solitary tract
and nucleus; N.m.X, motor tenth nerve nucleus (Vagus);
Tr. sp. V., descending trigeminal tract and nucleus; N. XII,
twelfth nerve; 01.i., inferior olive. (From Brodal, Note 80)

NEURAL SUBSTRATES OF ATTENTION MECIIANISMS 589

 •••••
•••••••••
tab •

• I • .•
1 • •

••
• 1 •

1•.

•
• -Bs
••••••••
••••

CM
FIGURE 11 Distribution of reticular cells sending axons rostrad (
left) and caudad (right) is indicated in two semi-schematic sagit-
tal sections of brain stem of cat. Although there is a good deal of
overlapping, caudally directed axons appear to arise somewhat
more rostral than rostrally directed fibers. The arrows at the side
• : ••• •• •
of the figures indicate that all axon systems are both crossed and -
*:
I s se.
•• • . • ".
uncrossed, except fibers descending from pons, which are un-
crossed. Abbreviations as in previous figures. (From Brodal,
• • •
• /I I'
as.7• MI
Note 80)
••• .. 8
.. • • • • N, •••:*.;.• ...
•■■■•••■• . ••• 8
*..... • OM • •• .. •
•• ••• ``- •••• • o• • ••
discriminate mingling together of rostral- and caudal- •••••••
•

projecting cells, or a source in common from many of the

same neurons (Figure 11). Our Golgi studies indicate that
both of these alternatives exist.
Anterograde studies, employing modifications of the
Bielschowsky reduced-silver method, allow certain state-
ments to be made as to terminal stations of ascending and
descending projections. Reticulo-spinal fibers have been
followed in ventral and lateral funiculi to termination in
spinal laminae 7 and 8.12 In some cases, Golgi preparations
of spinal cord show such fibers terminating along den-
drites and somata of internuncials and occasionally along
the outer segments of motoneuron dendrites. In the latter,
the anatomical material is not sufficiently precise to dif-
ferentiate alpha from gamma motoneurons."
Rostral projections have been followed by a number of
workers using Marchi, Glees-Bielschowsky, and Nauta
methods (Figure 12). There is general consensus that the
ascending system projects dorsally into the thalamic
intra-
FIGURE 12 Degeneration of ascending fiber systems, as seen in
sagittal section following a lesion in the caudal midbrain teg-
mentum, visualized by means of the Nauta method of antero-
grade degeneration. Large dots represent fibers of passage; fine
stipple represents probable preterminal axons. LM, medial lem-
niscus ; IP, interpeduncular nucleus; MI, massa intermedia ; CS,
superior colliculus; NR, nucleus ruber ; MT, mamillothalamic
tract; V, ventral nuclear complex; L.DL., lateral dorsal nucleus;
DM, dorsomedial nucleus; BC, brachium conjunctivum; SN,
substantia nigra; AD, anterior drosal nucleus; AM, anterior
medial nucleus; f, fornix; CA, anterior commissure; CP, pos-
terior commissure; PM, mamillary peduncle; CM, mamillary
body; Pc, paracentral nucleus; CI, inferior colliculus. (From
Nauta and Kuypers, Note 95)

590 BRAIN ORGANIZATION

laminar system, especially to the paracentral and central
lateral nuclei and possibly as far rostrally as the nucleus
reticularis thalami. Although there are some reasons—both
anatomical and physiological—for expecting terminations
in centre median, it has also been suggested that degenera-
tion granules found in this area are purely of en passage
variety and do not represent functional preterminal ele-
ments." The ventral prolongation of this system can be
traced into the zona incerta of hypothalamus and the fields
of Forel. Lesions placed progressively more rostrally and
ventrally in mesencephalic tegmentum reveal axonal sys-
tems that reach preoptic, septal, basal-forebrain, and basal-
ganglionar stations. The vexing problem of whether direct
monosynaptic connections are established with cortex by
any of these elements has not yet received a satisfying an-
swer from these techniques 9 5
As useful as these tracking methods have proved to be,
it remains for the Golgi methods to demonstrate the rich-
FIGURE 1 3 Sagittal section of an entire mouse brain (7 days
old) showing 2 reticular cells in the magnocellular nucleus
of rostral medulla. Both cells emit axons that bifurcate and
course rostrad and caudad. A number of collaterals are given
off by each axon, some of which reach cranial nerve nuclei,
such as Deiter's component of the vestibular complex, n.D.;
both inferior and superior colliculi, I.C. and S.C.; and pre-
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
ness and complexity of reticular projection systems—
preferably viewed in sagittal sections of postnatal mice and
rats (Figure 13). Although admittedly "simpler" than
carnivores and primates, the rodent brain stem can well
serve as a paradigm for higher forms, while escaping much
of the circuit redundancy that makes Golgi analysis of
larger brains so difficult.
The axons of reticular neurons form a spectrum of con-
ductors of many lengths. Some neurons appear to project
rostrally only; and others only caudally, but a significant
number of medium-sized and large reticular elements gen-
erate the familiar bifurcating axon (Figure 14) that may
project to distant stations both upstream and down. Vir-
tually all of these are characterized by collaterals of vary-
ing lengths that leave the main stem perpendicularly and
penetrate into adjacent reticular areas. Longer collaterals
may reach cranial nerve nuclei and/or sensory relay and
extrapyramidal motor fields. A number of fiber counts
tectum, Pt. Other abbreviations include CM, centre me-
dian; LP, lateral posterior; LG, lateral geniculate; LD,
lateral dorsal; CL, central lateral; AV, anterior ventral; V,
ventral complex; VA, ventral anterior; R , nucleus reticu-
laris thalami; ZI, zona incerta; and SN, substantia nigra. (
From Scheibel and Scheibel, Note 33)
591
 FIGURE 14 Sagittal section through the brain of a young
rat showing the axonal trajectory of a single neuron of the
nucleus reticularis giganto-cellularis, R. The rostral-coursing
axonal component supplies collaterals to inferior
colliculus, j ; the region of the III and IV nerve nuclei, i;
mesencephalic tegmentum, h; posterior nuclear complex of
thalamus, f; dorsal, intralaminar and ventral thalamic nuclei
respectively, e, d, and c; zona incerta of hypothalamus, g;
nucleus reticularis thalami, b; and basal forebrain area, a.
The posterior-
have suggested to us that, on the average, each reticular
axon releases one collateral of approximately 100-micron
length for each 100 microns of trajectory." This does not
include large numbers of smaller, bulb-like enlargements
along the course of the axon, called by Cajal boutons-en--
passage5 and representing, according to electron-micro-
graphic analysis, centers especially rich in mitochondria
and other intra-axonal organelles.
The picture that emerges is one of continuous, intensive
interaction between large numbers of conductors and the
surrounding matrix of core neurons. The nature of these
contacts must vary, depending on the presence or absence
of a myelin sheath and the frequency of axonal specializa-
tions. No definite statement can be made about the func-
tional significance of the untold numbers of contacts es-
tablished among structures without discernible axonal (or
postsynaptic) specialization. It is currently fashionable to
discount the significance of such membrane appositions as
seen in electron micrographs if there is no evidence of
membrane thickening, increased opacity to the electron
beam, or presence of synaptic vesicles behind one of the
592 BRAIN ORGANIZATION
directed component sends collaterals into the substance of
the reticular core, m; the hypoglossal nucleus (XII), k; the
nucleus gracilis, 1; and the intermediate gray matter of the
spinal cord, n. (This illustration was originally prepared for
publication in Brazier, M. A. B., "The Electrical Activity
of the Brain"; Third Edition, New York, The Macmillan
Company; in press, and is reproduced here with permission
of the author.)
apposed membranes. However, the presence of trans-
membranal electrogenic effects has been shown to exist at
least in invertebrates,96.97 and Nelson and Frank have
mapped a considerable field effect around spinal motoneu-
rons in the cat." Pending more rigorous analysis with re-
cording techniques of higher resolution, no definitive
statements can be made about the presence or absence of a
spectrum of field interactions and threshold manipulatory
effects in a dense complex of long conductors and axoden-
dritic neuropil such as the reticular core.
However, the patterns of connectivity that can be traced
out through the core suggest a circuit scheme so richly
redundant that convincing arguments could be advanced
for almost any conceivable loop or chain. Figure 15 sum-
marizes three alternatives. The first depends largely on
polysynaptic chains of short-axon elements similar to the
scheme originally suggested by Moruzzi and Magoun.22
The second illustrates the collateral-rich, long-projecting,
high-priority axon now known to characterize many re-
t i c u l a r
neurons. The third combines a group of these, geo-
graphically staggered, so that each is activated somewhat

FIGURE 15 Schematic showing three possible circuit paradigms
through the reticular core of the brain stem. a: the chaining of
short-axoned cells as suggested by Moruzzi and Magoun. b : a
single long-axoned, high-priority conductor reaching from lower
brain stem (left) at least as far as diencephalon (right) without
synaptic interruption. This resembles a type of reticular axon
actually found in large numbers in the core. c: an ensemble of
r. such long axons, whose collateral systems, playing upon adjacent
elements similar in nature, but geographically staggered, can re-
sult in a spectrum of conduction latencies and divergence of in-
: formation. (From Scheibel and Scheibel, Note 33)
later in time, resulting in appreciable degrees of spatial and
temporal dispersion. Such an array could also account for
the appearance of a family of response latencies over intra-
reticular conducting paths, as demonstrated by Adey,
Segundo, and Livingston."
The rostral course of axonal elements of medullo-pon-
tine and mesencephalic tegmental neurons has already
been mentioned in part. While the dorsal thalamic leaf
almost certainly projects no farther than the intralaminar
fields and nucleus reticularis thalami, the ventral leaf, ex-
tending through ventrolateral hypothalamus, in some
cases may be traced to, and through, the basal forebrain area
and ventral caudate-putamen. By using very thick sections (
200 micron) of perinatal mouse, we have been able to
trace a limited number of axons from brain-stem reticular
formation to the base of the anterior third to half of cortex.
We estimate that the number of such monosynaptic corti-
copetal conductors is relatively modest—perhaps of the
order of 5 to 10 per cent. Although we are still working on
this problem, it is of interest that Magni and Willis34 have
recently succeeded in antidromically activating small
numbers of micropipette-impaled, brain-stem reticular
neurons via cortical stimulation. They consider their data
consistent with the presence of some reticular elements
that project monsynaptically upon cortex.
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
The thalamic system
The nonspecific system of the thalamus differs in organiza-
tional pattern and functional role from the remainder of
the brain-stem reticular core. It is also partially isolated
from reticular structures of the lower two thirds of the
stem, so it seems appropriate to deal with it separately. Ac-
cording to Jasperm100 "the thalamic reticular system is
the dorsal limb of the cephalic end of the brain stem
reticular formation. . . . [It] is sometimes identified with the
unspecific thalamocortical projection system. This may
not be accurate, since unspecific projections may be only
one important property of the thalamic reticular system."
Generally included within its domain is the calyx of medial
and intralaminar nuclei that run forward from the centre
median-parafascicular field to surround the dorso-
medial complex and separate it from the ventro-lateral
fields just outside. Mpre rostrally, the system includes—at
least at a functional level—portions of antero-medial and
ventral-anterior nuclei and the nucleus reticularis thalami
(Figure 16). The group of small cell masses making up the
internal medullary lamina itself includes central-medial,
paracentral, central-lateral, and a group of more variable
midlinebridging nuclei, i.e. reuniens, rhomboidalis,
interanteromedialis. Previous descriptions of these areas
may be found in various studies,5,101,101 while adequate
summarizing statements of their functional capabilities have
been offered by Jasper100,105 More recent discussions of
problems inherent in the experimental analyses of the
system have also been provided.94,106
The present discussion is limited to data that have
emerged from our analysis of thalamic nonspecific sys-
tems, using Golgi methods. It is worth passing comment
that problems in visualization of the intricate neuropil of
this system were noted by Cajal 60 years ago5 and probably
have been largely responsible for the dearth of information
on structural substrates of this intriguing area. Our own
material is presently based on analysis of approximately
2000 rodents, cats, and a few primates, and can be consid-
ered introductory.
Expressed in terms of its constituent neuropil as revealed
by the Golgi chrome-silver, Golgi-Cox, and ancillary
techniques, the thalamic nonspecific system consists of a
group of fields stacked along the rostro-caudal axis, and
connected by a dense feltwork of axons of varying lengths
and diverse projections. We shall consider the centre me-
dian-parafascicular complex as the common caudal com-
ponent, despite rather compelling evidence that the centre
median is a relatively late phylogenetic acquisition and is
virtually absent in rodents. From this complex, axons pour
rostrally (Figure 17), fanning out both laterally and medial-
ly; the latter elements bridge midline somewhat more
593
 7 411 s4 3 2 9 2 3 4 S • 7 •

\\
\ k- Fr.

*N-cp

Hp
•

I• 4 3 2 3 4 $ • 1 • • • 7 • r 4 s 0! 3 4 f
\ ss
1 1 7 • 1 1

A,10\
Fr. 6.0
Fr. 7.5 vs • " s \
'
\

\
\
s\

\ % •

•
•\'` . \ \ ' ‘ \ \ \

GM

I H•

Ped; SN

FIGURE 16 The electrophysiologically determined limits of cleus ; mc, pars magnocellularis ; MD, medial dorsal nucleus;
the nonspecific (reticular) system of the thalamus. AD, an- MFB, medial forebrain bundle; NCM, central medial nu-
tero-dorsal nucleus; aHd, dorsal hypothalamic area; AL, cleus; NCP, posterior commissural nucleus; NR, red nu-
ansa lenticularis; AM, antero-medial nucleus; AV, antero- cleus; P, posterior nucleus; Pc, paracentral nucleus; Ped,
ventral nucleus; BC1, brachium of the inferior colliculus ; cerebral peduncle; Prt, praetectum; Pt, parataenial nucleus;
CC, corpus callosum; Cd, caudate nucleus; Cl, claustrum ; Pul, pulvinar ; PVA, anterior periventricular nucleus; PVH,
CL, central lateral nucleus ; CM, centre median nucleus ; CP, periventricular hypothalamic nucleus; R, reticular nucleus;
posterior coinmissure; Da, nucleus of Darkschewitsch; En, RE, nucleus reuniens ; S, medullary stria; SG, supragenicular
entopeduncular nucleus; Fil, filiform nucleus; fsc, subcallo- nucleus; Sm, submedian nucleus; SN, substantia nigra ; Spf,
sal fasciculus; FT, thalamic fasciculus; Fx, fornix; GC, cen- subparafascicular nucleus; ST, terminal stria; THP, habenu-
tral gray matter; GM, medial geniculate body; HbL, lateral lopeduncular tract; TMT, mammillothalamic tract; TO,
habenular nucleus; HbM, medial habenular nucleus; Hi, optic tract; 'FTC, central tegmental tract; VA, ventral an-
field of Fore1; HL, lateral hypothalamus; Hp, posterior hy- terior nucleus; VL, ventral lateral nucleus; VM, ventral me-
pothalamus; IAM, interanteromedial nucleus; IP, inter- dial nucleus; VPL, ventral posterolateral nucleus; VPM,
peduncular nucleus; Is, interstitial nucleus; LD, lateral ventral posteromedial nucleus; ZI, zona incerta. (Figure and
dorsal nucleus; Lim, nucleus limitans; LM, medial caption from Jasper, Note 100)
lemniscus ; LME, external medullary lamina; LP, lateral
posterior nu-

594 BRAIN ORGANIZATION

 FIGURE 17 Horizontal sagittal section through diencepha-
lon of 12-day-old mouse, showing certain aspects of rostral-
coursing axonal projections from the thalamic nonspecific
system and the brain stem reticular core. The latter, f. Rt.,
runs through the lateral aspect of the intralaminar nuclei,
giving off collaterals to, and lying in parallel with, the pro-
jection of the thalamic system that here is limited to a small
number of fibers pictured issuing from the centre median-
parafascicular, CM-Pf complex, although the amount of
centre median tissue present in the rodent is uncertain (see
text). Collateralization of these systems appears to be both
ipsilateral and contralateral, extending into specific and
nonspecific nuclear masses. Most of these fibers are shown
rostrally in the reuniens and rhomboidalis components of
the massa intermedia. Rostro-lateral components peel off
in arcs, flowing through or around more lateral nuclear
masses to reach putamen and caudate.
Although it has generally been agreed that no axons of
the posterior half of the thalamic nonspecific system reach
cortex, more sensitive tracking methods in the hands of
some investigators1136 now suggest that delicate cortical
projections may, in fact, exist. Golgi methods have not
enabled us to trace such axons to their destination, al-
though there seems no question about cortical termina-
tions for more rostrally situated elements. The Golgi
methods do, however, show the intensive collateralization
of these caudal and rostral components, which provide a
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
terminating no farther rostral than the nucleus reticularis
thalami (Retic.), although there is some evidence that ele-
ments among them may reach striatum and even cortex (
see text). One such fiber is shown at a, while LS. represent a
small group of specific thalamo-cortical elements. Other ab-
breviations include: Pt, parataenial nucleus; AV, anterior
ventral nucleus; AM, anterior medial nucleus; IAM, in-
teranteromedial nucleus; PC, paracentral nucleus; CeM,
central medial nucleus; CL, central lateral nucleus; IV, in-
terventricular nucleus; GL, lateral geniculate; GM, medial
geniculate; Rh., rhinencephalon. (From Scheibel and
Scheibel, Note 33)
rich substrate for interaction among the various fields of
the nonspecific system and with adjacent specific and as-
sociational nuclei. As suggested by Figure 17, the ascend-
ing axonal system from the posterior two-thirds of the
brain stem runs roughly parallel to the thalamic nonspe-
cific system and provides one of its most important sources
of afferent excitation.
This relationship is shown more clearly in F i g u r e 18,
where a dense collateral mass infiltrates the intralaminar
system from ascending brain-stem reticular fibers as well
as from the adjacent, much attenuated spino-thalamic
tract. Individual elements of this influx penetrate not only
the entire ipsilateral intralaminar system, but reach some
of the contralateral fields—at least in the rodent (specifical-
595
ly rats and mice). We have not yet been able to identify
the entire category of afferents to this system, nor is such
information available from degeneration and/or evoked-
potential studies. However, we have identified axonal ele-
ments from basal ganglia; the fields of Forel; colliculi and
pretectum; from adjacent associational nuclei such as lat-
eral posterior and pulvinar; hypothalamus; the anterior
nuclear complex; the stria medullaris-habenula-fascicu-
lus-retroflexus complex; and a massive, fine-fibered con-
tingent from (at least) orbito-frontal cortex. The nature of
this afferent array suggests that the thalamic nonspecific
fields are more sheltered from the onslaught of externally
derived data than is the reticular mosaic of the lower brain
stem.
This presynaptic influx reaches the thalamic fields along
FIGURE 18 Mass of collateral and terminal fibers from as-
cending reticular components (AR) and spino-thalamic
tract (Sp) terminate in the posterior half of the intralaminar
fields, including parafascicular (Pf); paracentral (Pc); central
lateral (CL); central medial (CeM), and project contralat-
596 BRAIN ORGANIZATION
two predominant axes, rostro-caudal and transverse. The
majority of dendritic elements of this system appear to be
organized transversely (see Figure 19); so we must assume
that those conductors with terminal elaborations in the
rostro-caudal axis either develop less potent synaptic drive
along the dendrites or terminate preferentially on somata.
Although the Golgi evidence is suggestive in this regard,
the data cannot yet be considered definitive.
Intensive pentration of adjacent, specific thalamic
fields is a common feature of this system. Figure 20
shows a number of such elements entering the neuropil mass
of the ventral-lateral (VL) nucleus, whose main afferent
supply comes from cerebellum via brachium
conjunctivum. Physiological support of this relationship
is supplied by Purpura, et al., who have found a
frequency-specific type
erally via the reuniens nucleus (Re). Other abbreviations in-
clude Aq, aqueduct of Sylvius ; Pv, periventricular fibers;
and fr, fasciculus retroflexus. Ten-day-old rat. Rapid Golgi
modification.
 FIGURE 1 9 Axonal and dendritic organization of portions
of the thalamic nonspecific system. Orientation of den-
drite masses of neurons in paracentral (Pc), central lateral (
C1), and anterior medial (Am) appears largely transverse or
oblique, thus paralleling contralateral axonal inputs, b, and
collaterals from descending centrifugal fibers, d. Some in-
tralaminar axons, like e, seem to remain largely ipsilateral,
connecting adjacent nuclei, while others, such as f, project
laterally into adjacent fields of the ventral nuclear complex
of modulatory control exerted by intralaminar nuclei
upon a background of ongoing, brachium-derived excita-
tion in VL neurons.107
In the rodent, massive commissural patterns are formed
by intralaminar axons. A series of fibro-nuclear masses
that enable intensive bilateral communication have been
described, including the nuclei reuniens, rhomboidalis,
and interantero medialis. This allows for a massive system
- intermedia bridging,'" there may reside substrate for the
of re-entrant loops of varying lengths' connecting both
sides of the thalamus, and also facilitates communication of
nuclear fields with both cortical hemispheres. The inten-
sity of such crosscommunication decreases as the phylo-
genetic scale is ascended and midline-bridging tissue
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
(see figure 20). A number of intralaminar-derived axons
such as c and d collateralize contralaterally and then project
rostrad toward striatum and/or cortex. Similar patterns are
found in axons from cells of anterior ventral (AV) and ven-
tral anterior (VA), which are not included in the intralami-
nar complex. The two axons marked a follow trajectories
of this type. Other abbreviations include Aq, aqueduct of
Sylvius ; and fr, fasciculus retroflexus. Twenty-day-old
mouse. Rapid Golgi modification.
shrinks in size and importance. One might wonder wheth-
er this feature of rodent thalamo-cortical anatomy might
not be substantially implicated in the phenomena of corti-
cal equipotentiality described by Lashley108 in rats (Chow,
this volume). In a similar vein, with the progressive de-
crease of commissural mass to the point at which 30 per
cent of the human thalami may be totally devoid of massa
rise of laterality in primate and man.
It appears highly likely that a cortically directed projec-
tion arises from the anterior third of thalamic nonspecific
fields. In Figure 21, part of this system can be seen leaving
the critical antero-medial angle of the field. Such axons
597
may project rostrally, unilaterally, or bilaterally, and as
they emerge from this pool, send rich collateral masses in-
to adjacent nuclei such as ventral-anterior and nucleus
reticularis thalami.110,111
The latter is of special interest to us. It continues as a nu-
clear plate or shell surrounding anterior, superior, and in-
ferior surfaces of the thalamus, and all thalamo-cortical
projections and the greater part of the cortico-thalamic re-
flux must penetrate through it. It is, accordingly, in a criti-
cal position to monitor and modulate most thalamo-cor-
tico-thalamic interactions. Several investigators111,112 have in
fact, likened it to the screen grid in a vacuum tube. Our
Golgi studies indicate that the dendritic ensemble of this
field is densely intertwined and covered with long, hair-
like spines, forming a net or sievelike structure through
which most axons of these thalamo-cortico-thalamic sys-
tems perforate, either directly or with production of col-
FIGURE 20 Vignette taken from a horizontal sagittal sec-
tion through the thalamus of a 7-day-old rat showing some
interrelations between thalamic nonspecific system fibers and
the ventral lateral and ventral anterior fields. Cells and fibers
of the thalamic nonspecific system (Thal. Retic.)
culminating in the rostral-running projection or
nonspecific radiation (nsr) send terminals and collaterals into
the neuropil fields of ventral lateral (VL) and ventral anterior (
VA) nuclei, whose primary afferent sources are the
dentato-rubro-thalamic,
598 BRAIN ORGANIZATION
laterals.111 Opportunities for intensive axodendritic inter-
actions arc obvious (Figure 22).
The axonal projection of the nucleus reticularis thalami
is clearly of great physiological importance. Cajal reported
that these axons seemed to run caudally,' but a long series
of retrograde degeneration studies following massive, se-
lective, cortical ablation appeared to favor a cortical ter-
mination.113,114 As a result of evoked-potential studies, it
was asserted by some investigators that the nucleus reticu-
laris thalami actually constituted the final common path-
way along an intralaminar polysynaptic chain to cortex.21
The importance of such data was somewhat weakened
by the inability of the stimulating electrode to differen-
tiate between nucleus reticularis cells and fibers of passage.
Recent Golgi studies have shown clearly that at least 95 per
cent of the axons of nucleus reticularis thalami cells are, in
fact, distributed caudally to thalamus and mesencephalic
(drt) and pallido-thalamic (pt) bundles respectively. Also
visible are sectors a, b, c, and d, of VA, the nucleus reticu-
laris thalami (nR) more anteriorly, and the ventrobasal com-
plex (VB) more posteriorly, receiving the terminating me-
dial lemniscus (lem). One centrifugal fiber (ct) is seen ending
in VB, while the ascending reticular system (ars) sends col-
laterals toward the VA field. Rapid Golgi modification. (
From Scheibel and Scheibel, Note 110)
 FIGURE 21 Horizontal section through the anterior end of
the thalamus showing the area of origination of the thalamic
nonspecific projection upon cortex. Collaterals and termi-
t nals from the ipsilateral nonspecific radiation, nsr (ips), and
some elements from the contralateral system, nsr (con) fill
the medial portion—sector a of VA. This region is also
reached by individual elements of the pallido-thalamic bun-
dle (pt), which arborizes maximally in sector c, while shorter
collaterals from the ascending reticular system (ars) termi-
nate in sector d. One rostrally-directed axon (vb1) from a
tegmentum (Figure 23), thereby providing a hierarchy of
re entrant circuits." Such feedback loops appear to play
- cends that produced by stimulation of specific thalamic
on neurons of both specific and nonspecific thalamic sys-
tems and may serve as substrate for recruitment poten-
, tials.18,19 They may also provide circuitry essential to the
alternating excitatory postsynaptic potential (EPSP) and
inhibitory postsynaptic potential (IPSP) swings described
in specific thalamic cells secondary to afferent barrage."
TIIALAMO-CORTICAL PROJECTION The projection of
thalamic nonspecific systems upon cortex poses a problem
that requires further investigation. Investigators do not
fully agree on how recruitment potentials are distributed
over cortex following intralaminar stimulation," ,2° al-
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
ventrobasal cell (VB) is seen, as are 3 heavy caliber and 2
fine cortico-thalamic fibers (et 1-5), the first two of which
terminate in VA. Other abbreviations including those in the
inset diagram include: 1, parataenial ; 2, anterior ventral,
AV; 3, anterior medial, AM; 4, interantero-medial, IAM;
5, paracentral, PC; 6, central lateral nucleus, CL; 7, centre
median-parafascicular complex; 8, interventricular ; C Str,
corpus striatum; Rh, Rhinencephalon. Three-day-old kit-
ten. Rapid Golgi modification. (From Scheibel and Scheibel,
Note 110)
though there is a consensus that the distribution trans-
nuclei. Our own investigation of this projection is still in-
complete, but the data are suggestive. In rats and mice, the
axonal outflow from the anterior third of the intralaminar
fields appears to project rostrally and ventrally via the in-
ferior thalamic radiation, just lateral to the mass of the
septal nuclei. The fibers can be followed forward to ap-
parent simple ascending (axodendrite?) cortical termina-
tions in orbito-frontal cortex. Each fiber may subdivide
into many branches while still in the subgriseal white mat-
ter, and the most caudal branches may reach almost to the
frontal motor fields.
Such data appear to complement the findings of Velasco
599
and Lindsley,116 who observed that orbito-frontal ablation
in the cat effectively blocks recruitment phenomena over
the remainder of cortex, thereby identifying orbito-fron-
tal fields as primary cortical distribution stations for as-
cending intralaminar control. On the other hand, Purpura
has removed large blocks of frontal cortex and underlying
subcortical tissue without disturbing recruitment phenom-
ena117 If further structural and functional studies should
bear out the importance of the orbito-frontal field as a dis-
tribution center, it would be logical to suppose that the
further backward spread of recruitment waves over cortex
would depend on a complex series of cortical chains of
varying lengths.
FIGURE 22 Portion of the anterior ventral sector of the nu-
cleus reticularis thalami in 12-day cat as seen in sagittal sec-
tion. The dense matting of the dendrite mass is seen paritcu-
larly along the inner border of the nucleus, nR; a, initial
intranuclear path of an axon with its primary collateral sys-
tem limited to the n. reticularis; al, its caudal path; a2,
penetrating the ventral anterior nucleus, VA; and a typical
bushy terminal dendrite structure, a3. Fibers b and c, are
centrifugal and centripetal elements, respectively, pene-
trating the nucleus and emitting collaterals b' and el en
600 BRAIN ORGANIZATION
Conclusions
Disparate but converging bodies of evidence now enable
us to define certain characteristics of the reticular core of
the brain. It is an archaic, centrally located mosaic of sub-
centers almost completely shielded from direct contact
with the self—world interface, yet continuously apprised
of all transactions crossing that interface. Its outputs, con-
tinuously variable along an intensity continuum, do not
reflect stimulus mode so much as stimulus quantity. This out-
put is projected simultaneously upon downstream effector
centers, upon rostral mechanisms devoted to differen-
tiation and comparation of information, upon primary
passage. The entire observed course of a fiber of unknown
Drigin, d, lies within the nucleus. Two cells, e and f, of the
ventral anterior nucleus, show dendritic configuration typi-
:al of this field. Axon g is emitted from a n. reticularis cell
whose soma remains unstained. Cell h of the n. reticularis
emits dendrite h', projects into more rostral white matter
cerebral peduncle), and loses its filamentous spines as it
does ;o. Other abbreviations include VB, ventrobasal
complex; Col. s.; superior colliculus ; Teg., Tegmentum;
Caud., caudlate. (From Scheibel and Scheibel, Note 111)

FIGURE 2 3 Slightly schematized horizontal sagittal section
through the thalamus of the mouse, showing some thalamic
afferent and efferent constituents on the right, and the course
of two nucleus reticularis thalami axons on the left,
elements h and i and two n. reticularis neurons. At h4 and h5
the axon of h collateralizes within n. reticularis, while h1,2,3
represent collaterals generated along the length of the axon
in its cau-
and secondary fields that relay raw information rostrally,
and upon the remainder of the reticular core itself. Its vol-
ume is small in comparison with cortex and cerebellum,
and economy probably demands repeated use of its rela-
tively limited amount of modular logic.62
It seems likely that stimulus patterns continuously re-
circulate through the millions of re-entrant loops as de-
cisional modes are reached. At the same time, competition
for the interest of the reticular arrays must be high, and
neural supremacy is gained for that moment in time only
NEURAL SUBSTRATES OF ATTENTION MECHANISMS
Teg. Mes.
dal trajectory and projecting into specific and nonspecific
thalamic fields. Similarly, in the case of neuron i, structures
marked i2,3,4 represent collaterals infiltrating the thalamic
ventral and lateral nuclear masses while it represents a short
collateral projecting rostrad into the striatum (Str). All other
abbreviations as in previous figures. (From Scheibel and
Scheibel, Note 111)
by those data that are most "exotic"—or most compelling
biologically. Like some stern, harried father figure, the
core has limited patience and limited time-binding re-
sources. Its logic is wide but superficial, and its decisionary
apparatus does not permit the luxury of hesitation.
Summary
Modern structural and operational concepts of the reticu-
lar core of the brain stem have emerged over the past 60
601
years, although a number of current ideas were already
implicit in studies published at the turn of the century. The
complex may be thought of as a mosaic of subcenters
placed athwart all input and output systems of the neu-
raxis, receiving a constant sampling of information from
activity ongoing in these systems. This mass of heterogene-
ous, convergent data is integrated along the postsynaptic
membrane of individual reticular neurons whose output
represents an algebraic summation of these inputs. The
output of these reticular-core elements, representing in-
tensity rather than mode, is then projected rostrally and
caudally to modulate the degree of synaptic drive on neu-
Subcortical and Cortical Mechanisms
in Arousal and Emotional Behavior
ALBERTO ZANCHETTI
THE DIFFERENT EMPHASIS that has been placed at different
times upon neural activity at either the cortical or the sub-
cortical level can be well exemplified by the history of re-
cent theories of emotion. Psychological thinking on this
subject was dominated at the end of the last century and
throughout the first three decades of the present one by
the so-called James-Lange theory.1,2 The neural basis for
this theory is schematized in the drawing of Figure 1 (left),
taken from a paper by Cannon.3 As Cannon says, "accord-
ing to the Jarries-Lange theory an object stimulates one or
more receptors (R, in Figure 1), afferent impulses pass to
the cortex (path 1) and the object is perceived; thereupon
currents run down to muscles and viscera (path 2) and al-
ter them in complex ways; afferent impulses course back
to the cortex (paths 3 and 4), and when there perceived
transform the 'object-simply-apprehended' to the 'object-
emotionally-felt' ; 'the feeling of the bodily changes as
they occur is the emotion—the common sensational, asso-
ciational and motor elements explain all,' to quote James's
expression." Obviously, then, in the James-Lange theory
ALBERTO ZANCHETTI Istituto di Patologia Medica, Univer-
sity di Milano, and Gruppo Nazionale di Medicina Sperimentale
del Consiglio Nazionale delle Ricerche, Milano, Italy
602 BRAIN ORGANIZATION
rons spread widely throughout cortex, brain stem, spinal
cord and, in some cases, as far peripherally as the first-
order sensory cell.
An impressive group of physiological mechanisms arc
subserved by this structural substrate and include gating of
sensory inputs; modulation of transmission along major
rostral-coursing and caudal-coursing tracts; modulation
and control of spinal effector mechanisms; multileveled
control over most visceral functions; and the active manip-
ulation of a spectrum of states of consciousness from deep
coma through a series of sleeping states to maximal vigi-
lance.
the cerebral cortex plays a predominant role in emotion,
first in simply apprehending the object, then in feeling it
emotionally. If we want to state this somewhat differently,
the cerebral cortex enjoys predominant attention in the
James-Lange theory because it refers exclusively to emo-
tional feeling.
Cannon3,4 strongly objected to James's and Lange's
opinions and formulated the so-called thalamic theory of
emotion. The neural basis for this theory was schematized
by Cannon3 himself in the drawing reproduced in Figure
1 (right), and has been conveniently summarized by
Lindsley5 as follows: "An external emotion-provoking
stimulus excites receptors (R) and starts impulses toward
the thalamus (path 1). . . . Thus efferent discharges are set
up in path 2, either through direct activation of the thala-
mus over path 1 or after impulses have passed to cortex (
path 1'), where they inactivate inhibition over path 3,
which allows patterned motor responses in the diencepha-
lon to find expression in effectors via.path 2. At the same
time an upward discharge in path 4 carries to the cortex
an appreciation of the pattern just released. . . . The differ-
ence between this view and that of James and Lange is that
for Cannon 'the peculiar quality of the emotion is added
to simple sensation when the thalamic processes are
roused.'
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