cns Flashcards
CNS basics
CNS = brain + spinal cord
* protected by bony structures (vertebrae) + 3 layers membranes = meninges
* receive afferent from PNS, analyse, sompare w past, + integrate to gen motor output to PNS
brain regions
forebrain = cerebrum + diencephalon w pit gland, limbic sys + olfactory bulb
midbrain = midbrain
hindbrain = pons + medulla oblongata + cerebellum
cerebrum has longitudinal fissure => L + R hemispheres
spinal cord
out brain @ foramen magnum, down vertebral canal
* each spinal seg gives off pair segmental spinal nerves to supply diff region of bod
* at caudal end canal cord peters out + spinal nerves for caudal structures form cauda equine
timeline brain development
- dorsal ectoderm thickens form neural plate
- other development processes overlap + continue well into adulthood
neurulation
- notochord = cylinder mesodermal cells + secr chem signals inc prot sonic hedgehog induce formation neural plate
- chem signals also stim other mesodermal cells develop -> vertebral bods + intervertebral discs of spinal column (form protective structures)
- plate indented along midline = neural groove
- groove deepens + ectodermal walls thicken @ dorsal lips = neural folds
- folds fuse to close tube, starting cervical region + moving both rostral + caudal
- rostral/caudal neuropore = pt closed to (hole) - get smaller + smaller + close last
- dorsally either side superficial ectoderm proliferates -> cells -> pinch off + form neural crest
which parts from neurulation form what
CNS from neural tube
PNS from tube + neural crest
what happens after neurulation
rostral neural tube forms vesicles bc its hollow
* first 3 primary vesicles develop
* these differentiate into secondary vesicles
* these develop to adult brain structures
which mols control patterning which parts CNS
- patterning ventral structures controlled by sonic hedgehog prot from notochord
- patterning dorsal structures controlled morphogenetic prot signals frojm superficial ectoderm
names of diff vesicle structures + development path
all early in gestation
differential growth means diff outpockets form
brain flexures at vesicle stage
important to orient structures
cerebrospinal fluid is + why
bathes inside (w/in tube) + out (its in meninges) of CNS for nourishment, waste removal, protect brain
spina bifida occulta
incomplete rostral or caudal fusion of neural folds = tube open = vertebral arches can’t form + fuse = neural v close outside world
* caudally common tailless breeds
grey matter + white matter
grey = cell bodies, dendrites + synapses
white = axons - white bc of fat in myelin
glial cells
supporting cells in CNS
nuclei + tracts w/in contect CNS
nuclei = clusters of cell bodies in CNS (ganglia in PNS)
tracts = bundles of axins in CNS (nerves in PNS)
decussation of tracts
many CNS functions cross over bet 2 hemispheres cerebrum
== info from/output to R side bod handled by L side brain
cross section spinal cord vs cerebrum
spinal cord = grey matter inside, white out
cerebrum = white + basal nuclei in deep parts w grey formed over
corpus callosum
white matter tract connecting cerebrum hemispheres, allowing exchange info
corpus striatum
basal nuclei in cerebrum interwoven w white matter tracts = stripy appearance
* involved planning + executing (esp habitual) movements (motor function)
* input from motor cortex + integrate w other inputs (thalamus, limbic sys)
* important animals, e.g. birds, w/o developed cerebral cortex (instead of motor cortex)
lining neural tube
initially pseudostratified columnar then diffs 3 layers:
1. inner ventricular zone = germinal = ependymal = gen new cells (divide into)
2. mantle = neuroblasts migrated here + diff into neurones/glia (+ mitotic in interphase)
3. marginal = axons of neurones w cell bods in mantle = white matter
diagram cell types lining neural tube life cycle
lining -> ciliated to move CSF
surface cerebellum compared cerebrum
cerebellum = smaller folds tiss called folia (as opposed sulci/gyri)
important sulci/gyri to remember
- cruciate sulcus crosses longitudinal fissure at right angle w motor cortex controlling movement sat there
- sylvian gyrus involved in processing auditory info (looks like ear)
brain lobes
named after bone section sits under (ish)
what does frontal lobe control
- thinking
- speaking
- memory
- movement
== personality
what does parietal lobe do
- language
- touch
- taste
- smell
what does occipital lobe do
- vision
- colour
- letters
- L/R
what does temporal lobe do
- hearing
- learning
- feelings
- fear
cortexes: where + what
- motor cortex just rostral cruciate sulcus = conscious movement
- somato-sensory cortex just caudal = conscious sensation
- visual = caudal occ lobe (sight)
- auditory = ventral to cruciate sulcus, dorsal temp lobe (hearing)
- association cortex = rest, linking together, e.g. memory
developing cerebrum
- internal cavity telencephalic forms lateral ventricles
- cell bods form from mantle layer
- SHH + BMP drive arrangement of regions
interconnected ventricles comm w central canal (remnants central cavity) in spinal cord
lat ventr comms rest of tube via 4th ventr
how are brain ventricles connected
lat ventr in each hemisphere, CSF flows thru interventricukar foramen in to 3rd ventr, thru mesencephalic aqueduct -> 4th ventr -> central canal
CSF is circulating around interconnected spaces then down canal
holoprosencephaly
failure in patterning vesicles as prosencephalon (forebrain) fails divide = single-lobed brain + single central eye
* can be caused by corn lily ingestion (cyclopamine toxin) - interferes w SHH signal recog
rhinencephalon is?
= olfactory bulb + olfactory tracts + olfactory peduncle + piriform lobe + hippocampus + fornix ==> olfaction for survival
* comms w higher centres affect emotion, behaviour, comms
* size reflects importance olfaction to species
how does rhinencephalon work
- olfactory bulb receives nerve fibres of olfactory nerve as run thru nasal cavity (via cribriform plate)
- down olfactory tracts
- **piriform lobe ** is part cerebral cortex where olfactory info processed
development brain stem general
midbrain from mesencephalon
pons from metencephalon
medulla oblongata from myelencephalon
mantle layers diff parts organise into dorsolateral alar plates + ventrolateral basal plates
* sepped by central roof + floor plates (no cont neuroblasts)
development myelencephalon/metencephalon
- alar mantle lamina stretches + move laterally over roof (made ependymal cells) to flatten
- form rhimbic lips that join in middle form cerebellum
- thickening develops ventrally form pons + medulla oblongata
cerebellum structure
- central vermis
- 3 peduncles either side connects it to brainstem
- v folded cortex (-> folia) w central nuclei = grey matter
- white matter branches = arbor vitae
cerebrllum development
2 lateral hemispheres formed from rhombic lips metencephalon
* joining of lips forms narrow central vermis
what is frontal cortex involved w
- decision-making
- planning
- motivation
- judgement
pre-frontal cortex
most rostral cerebrum
* personality + social behaviour
what does cerebral cortex do
allows conscious perception sensory input, movement muscs + conscious thought
limbic sys
series tracts + nuclei deep w/in cerebrum key to emotion, learning + memory
* species driven by instincts w less control over emotions + behaviours have relatively larger
thalamus
sensory info to brain synapses here b4 going -> cerebral cortex (except olfactory)
* also involved sleep + wakefulness
hypothalamus
connects endocrine + nervous sys + forms part limbic sys (= role in behaviour)
function brain stem
- cont key CV, resp, GI control centres
- cont nuclei for cranial nerves except I + II
- route comms bet higher structures + spinal cord
- tectum in midbrain involved orienting head + body in response sights + sounds
cerebellum functions
- organises + refines motor activity
- coordinates gait, control musc tone + voluntary musc activity
- compares intended movement w outcome = integrating sensory inputs
- balance + coordination - dysfunction = loss inhibitory component = loss coord
cant initiate musc contraction, can only refine
how does degree folding cerebral cortex vary
varies bet species mainly depending on how much space available in skull (also correlation w intelligence but less important)
how does association cortex vary bet species
amount incr w complexity of species + degree to which use prior experiences + memories to govern functions
layers meninges
- dura mater (outside)
- arachnoid mater
- pia mater
functions of meninges
- physical protection central nervous tiss
- facilitate flow cerebrospinal fluid (CSF)
- provide framework for bvs supplying CN tiss
dura mater
- thick fibrous layer made collagen + elastic fibres
- only CN tiss that’s pain sensitive (trigem innerv)
- own blood supply
2 layers: outer periosteal + inner meningeal - usually indistinguishable except seps as goes into 2 fissures, forming venous sinuses:
1. falx cerebri in longitudinal fissure
2. tentorium cerebelli in transverse fissure
fused to endosteum
where is transverse fissure
bet cerebellum + cerebral hemispheres
arachnoid mater
thin transparent mem w fibres forming web structure in subarachnoid space
* avascular, v lil innerv
subdural space
potential space = dura + arachnoid mater v closely associated, not much space, just a lil lymph-like fluid
subarachnoid space
true space, criss-crossed by fine collagen trabeculae + filaments (connect mems)
* conts CSF + bvs (supply nervous tiss)
pia mater
thin mem
* v vascular w dense innerv
* v adherent underlying tiss to nurture
* follows bvs into tiss + merges w tunica adventitia
denticulate ligaments
given off by pia mater, extend across subarach space + arach mater to join w dura mater
* act as sling to support spinal cord so no vibrate, bounce (= protect it)
how do meninges develop
from mesenchymal tiss around meninges
1. axial mesoderm -> mesenchyme -> ectomeninx -> dura mater (= pachymeninx)
2. neural crest cells from ectoderm -> mesenchyme -> endomeninx -> arach + pia mater (= leptomeninges)
3. spaces in mesenchyme coalesce -> subarach space
epidural space
dura seps from vertebrae w/in vertebral canal, creating epidural space
* -> filled w fat + CT = extra protection
=> dura mater only of spinal cord only attached to bone at foramen magnum + caudal end
filum terminale
beyond end spinal cord dura mater narrows form cord-like ligament that inserts on vertebra
cuff zones
formed from meninges around roots cranial + spinal nerves
* => grape-like structures where arach coming to end
dura mater most perm drugs, arach resistant but arach ending = more perm so these regions allow drugs to more easily enter CNS
cererbrospinal fluid (CSF) composition
- cell-free
- prot-free
- low aas (bc they function as NTs in CNS)
- low stable K+ (or would affect elec activity neurones)
- lower gluc than plasma
function CSF
- physical protection CNS tiss
- circulate nutrients + NTs to CNS tiss
- make stable environ for neurones so random a pots no triggered
- vol buffer - can decr to accomodate small CNS swelling
all works bc comms w ISF of neural tiss freely thru pia mater + ventr lining
how keep CSF stable
caps in CNS low permeability = blood brain barrier (BBB)
* tight junctions bet endothelial cells
* thick basement mem
* specialised connecting glial cells (astrocytes) have foot processes that decr permeability
where find CSF
circs subarach space, brain ventrs + central spinal canal
* constant prod + drainage = press grad, helps circ
* ventr + canal lined ependymal cells w cilia = help circ + joined loose desmosones = CSF can exchange w ISF of neural tiss
where do ventricles develop from
- laterals = telencephalic vesicles
- 3rd = central cavity of telencephalon + diencephalon
- mesencephalic aqueduct = mesencephalic vesicle
- 4th = rhombencephalon
where is CSF proded
mostly choroid plexi + small amount = ultrafiltrate pial bvs
1. bet lateral + 3rd ventr, pumping into ventr
2. in roof 4th - pumping into ventr + subarachnoid space
how is CSF proded
- transporting ependymal cells covering choroid plexi pump solutes (inc Na+) -> CSF
- draws water in by osmosis
- lipid soluble substances pass readily into CSF, e.g. O2, CO2
- cells + large mols like aas can’t pass across
- water soluble mats have be actively transported in, e.g. gluc
varies abt drugs passing in - some diseases affect perm of BBB
where does CSF drain
- out small holes in 4th ventr = bilateral apertures -> subarach space
- arach mat extensions arachnoid granulation/villi push thru dura into sinus -> venous circ
also via veins + lymph vessels around roots spinal nerves
press grad = flow 1 way, but can’t flow other even if press grad flipped
how does pia mater adhere underlying tiss
- choroid plexi
- reticular + elastic fibres
- cytoplasmic process of astrocytes in neural tiss
what is choroid plexus
tiss tufts v vascular w network bvs in covered transporting ependymal cells in ventr
how do choroid plexi develop
- grooves form in ventromedial cerebral hemispheres = choroid fissures + pia mater covering it invags into lat ventrs
- tela choroidea = region pia mater that adheres underlying ependyma (neuroepithelial lining) invaginates in fold into roof 4th ventr in myelencephalon + trapped
how sample CSF
from subarach space:
1. where enlarged form cerebellomedullary cistern @ atlanto-occipital junction
2. where widens beyond end spinal cord form lumbar cistern (L5-6 space)
via cisternal puncture
atlas
C1 bone
why sample CSF
give idea of health - does it have prots in, less gluc than plasma etc
arterial supply CNS
centred round cerebral arterial circle on ventral aspect surrounding hypothalamus
* basilar + internal carotid arteries supply circle
* rostral/middle/caudal cerebral + rostral/caudal cerebellar arteries move to cerebral hemispheres + cerebellum
forebrain receives more than other areas
parenchyma
functional tiss of organ (distinguished from CT + supporting tiss)
possible arterial sources to circle
combo internal carotid, basilar, maxillary + vertebral
* varies bet species - important for humane slaughter to lose lots blood + lose consciousness quick = minimal pain (position of animal has effect)
circle arterial supply horse/dog
from internal carotid + basilar arteries
circle arterial supply cow
maxillary + vertebral artery via 2 retia (net small vessels)
* decr press b4 brain + blood cooled down
circle arterial supply sheep/cat
mainly maxillary artery via rete mirabile
* internal carotids obliterated after birth
* basilar flows caudal so only blood -> caudal medulla oblong
collateral circ in brain
v few interarterial anastomoses = v lil collat circ = lots functional end arteries = occlusion/rupture => ischaemia + infarction of tiss supplies
brain venous drainage
low press sys = accoms changes in blod vol to maintain preload
slow, bidirectional, intermittent bloodflow w thin walls = potential seeding tumour or infection
variation bet species
what part does dorsal sagittal sinus drain
veins of cerebral hemispheres (dorsal part brain)
what part do cavernous sinuses drain
rostral forebrain, nasal cavity, orbit face = easy for infection -> blood supply
internal carotids traverse sinuses = cool blood in species w/o rete
spinal cord blood supply
branches aorta (caudal) + vertebral artery (cranial)
aorta enters thru intervertebral foramina -> 2 branches
1. small dorsal branch
2. ventral spinal art from coalescing large ventral branches
venous drainage spinal cord
bilateral venous plexi in epidural space w loads anastomoses -> segemntal vertebral veins at intervertebral foramina
-> vertebral vein/azygous vein/cranial caudal vena cava depending where in bod
eye =?
- organ of sight - turns light -> a pots in retina (so need light -> there unhindered), optic nerve takes a pots -> visual cortex
- develops from optic cup of embryonic diencephalon
- in bony orbit surrounding supporting soft tiss structures (= adnexa)
eyeball layers
- outer fibrous sclera
- middle vascular uvea
- inner neural retina
shape layers maintained by internal support from humours
eye anatomy
canine/feline
aqueous humour
- transparent, colourless, water-like ~ low-prot plasma
- proded ciliary process (constant w constant drainage)
- maintains intraocular press (= HP) to maintain spherical eyeball
- provides nutrients to avascular structures (cornea + lens)
vitreous humour
transparent, colourless hydrogel to maintain retina against choroid (support globe) + provide nutrients
* lamellar arrangement prot fibrils maintains transparency (trap hyaluronic acid)
* cont phagocytes to remove unwanted cellular debris
* cont hyalocytes to turn over hyaluronic acid
* attached caudal lens capsule, ciliary body + periphery optic disc
same over lifetime so old = more fluidy, affecting vision
fibrous tunic
opaque sclera caudally + transparent cornea rostrally
* junction bet them = limbus = drainage pt for aqueous humour of anterior chamber
* support + structure globe
cornea general
rostral 1/4, avascular
* transmits + refracts light = has be transparent + smooth (also nutrition)
* v sensitive w nerve endings - opthalmic branch trigem => corneal reflex (touch + blink)
layers cornea
- epithelium (squamous outer -> columnar) - protect stroma + decr water entry
- Bowman’s mem - thick, organised collagen fibres keep transparent, supporting keratocytes
- stroma
- Descemet’s mem - proded endothel to sep from stroma, thicker as older bc prod constant
- endothelium - actively pumps water from stroma to maintain transparent (can’t replicate = thins as cells spread to cover damage)
fluorescein stains stroma, not Descemet’s mem - know level of ulcer
sclera
dense fibrous CT + elastic fibres
* protects internal eye structures + maintains globe shape
* vascular w attachment extrinsic muscs
* axons from retina pass thru at lamina cribrosa to form optic nerve
dull white colour
uvea
== choroid (posterior uvea) + ciliary bod + iris (anterior) + suspensory ligament
vascular + pigmented to stop light out back eye
* firmly attached sclera at exit of optic nerve, less firm elsewhere
iris
sphincter w dilator + constrictor muscs to alter size pupil (= opening in centre)
* radial + circular sm musc fibres
* pigmented cells give range colours (lack pigment = light blue)
* ANS occulomotor n. - symp dil (= mydriasis), para constr (= miosis)
* seps anterior + posterior chambers
regs amount light entering eye
rostral continuation ciliary bod
choroid
eye lining (all of uvea behind lens, 2/3)
* vascular: bvs to all internal eyeball structures
* darkly pigmented to prevent light rays escaping out back eyeball
bet sclera + retina
tapetum lucidum
triangular yellow-green iridescent area light-reflecting cells w/in choroid dorsal to where optic nerve leaves
* reflects light back to photoreceptor cells = pass thru again = improve night vision
well developed carnis, present most mammals but humans + pigs
ciliary body
thickened continuation of choroid
1. ciliary musc = sm fibres to control thickness + shape lens
2. ciliary process secr aqueous humour into posterior chamber
also anchors lens
pathway outflow aqueous humour
posterior chamber, thru pupil + drained @ iridocorneal angle = venous plexus so humour can return circ
-> via trabecular meshwork -> Schlemm’s canal -> episcleral veins
iridocorneal angle
junction bet corneal limbus, root iris + anterior ciliary bod
glaucoma
iridocorneal angle blocked = drainage stopped = intraocular press builds up = red eye
serious
suspensory ligament of uvea
continuation of ciliary bod, forming circular support round lens perimeter (zonular fibres)
* connect bod (musc) to lens, all around lens’ circumference
lens
transparent biconcave disc sepping aqueous + vitreous compartments
* suspended by ciliary bod
* cont outer capsule, regular arrangement lens fibres (from around equator to meet anteriorly + posteriorly) + central nucleus
* concentric layers allow eye to focus
contr/relax ciliary musc w suspensory ligs alter shape + change depth of focus = lens accomodation
distance vs close focus
distance = relax musc, tight lig = lens stretched, longer, thinner, less refraction light
close = contr musc, slack lig = lens thicker + refracts light more
cataract
lens opacity due disruption arrangement lens fibres
retina
innermost eye layer of caudal wall
1. light focused onto photoreceptors (light sensitive) by lens
2. converts light -> a pots -> brain by optic nerve
nutrition from choroidal bvs externally + retinal bvs internally (run from optic disc)
layers retina
from closest to choroid (where light hits last)
1. retinal pigment epithelium (no pigment over tapetum lucidum)
2. photoreceptor cells (rods + cones)
3. bipolar neurons
4. multipolar ganglion cells
pigmented layer retina
prevent light leaking out eyeball, augmenting effect pigment cells in choroid (work together)
photoreceptor cells
- rods - sensitive low light levels but not colour = black + white + night vision
- cones - sensitive bright light, provide colour vision
dogs + cats = 95% cones = light + shade but colour poorly developed
bipolar neurons of retina
gather info from rods/cones + transmit to next layer
ganglion cells of retina
axons run across surface retina -> leave at optic disc form optic n.
* unmyelinated so light thru -> photoreceptors
* at optic nerve attachment no rods or cones = blind spot
ocular fundus
structures of posterior eye visible on opthalmic exam
* optic disc
* retina w pigmented epithelium
* retinal bvs
* choroid inc tapetum - where visible = tapetal fundus (rest = non-tapetal fundus)
* sometimes sclera
how are dog + cat fundus diff
dogs = white bc nerves becoming myelinated b4 leave eye
cats = no myelinated until leave eye => black
how is rabbit fundus diff
bvs only medial + lateral + band not circle of myelination
how is horse fundus diff
dots of vessels thru choroid layer (= stars of Winslow)
* abnormal other species that have large clear retinal bvs from optic disc
how does field of vision vary
- predatory species, e.g. cat = eyes pt forward, wide area binocular vision (fields of eyes overlap) = pinpt position accurately
- prey = eyes side head, mostly monocular/uniocular vision - less detail but greater field view to be able see where is to run away
cilia on eyelids
angled away from eye for further protection entry foreign mat
resemble human eyelashes
palpebral ligaments
give palpebral fissure its shape + hold canthi in place
* medial + lateral ligs
conjunctiva
v vascular mucous mem lining inside eyelid front eye except cornea
1. on eyelid = palpebral
2. on globe = bulbar
- lots lymphoid tiss to protect against infection
- conts goblet cells prod mucin component tear film (stabilise it)
covers nictitating mem
tears
keep corneal surface moist bc dry = cornea damaged = loss vision; cont:
1. mucin stabilise tears + keep tear film on eye surface (de goblet cells conjunctiva)
2. lipid reduce evap tears (de Meibomian/tarsal glands)
3. aqueous component = hydrating for eye health (de lacrimal gland + gland of 3rd eyelid @ base)
Meibomian glands
== tarsal glands
* sebaceous, in eyelids, proding oily (lipid) secr so tear film no evap from cornea
Harderian gland
under 3rd eyelid, proding 1/3 aqueous component tear film
* cherry eye = enlarged + protrudes from under nic mem (brachycephalics common) - surgery push back behind
cranial nerves to eye
- optic = sensory, transmit visual info retina -> brain
- trigem - mixed, inc sensory to structures surrounding eye + intraocular surfaces
diffs bet species w eyelid
cat = eyelids well atached to outside eye (narrow palperal fissure)
rabbit = structures looser = less good at moving tear film over eye
horse = prominent lacrimal caruncle (corner eye w oil/sweat glands)
lacrimal puncta
opening that pumps tears out eyes
* most have dorsal + ventral, rabbit just ventral
species diffs w cornea
- ungulates = horizontal oval shaped
- rabbits retain ability replicate corneal endothelium into adulthood
species diffs iris + pupil
cat constr = slit, dil = circle
dog/pig = circular pupil
ungular = horizontal oval shape for max. horizontal field vision (constricts on horizontal plane)
where do optic nerve + retina develop from
outgrowth of wall of wall of diencephalon (optic cup)
fovea
notch in retina w bipolar + ganglion cells displaced so easier for light thru
* 1:1 synaptic contact
* area greatest visual clarity
* only cones then rods round outside of eye
photopigment in rods
rhodopsin
* converts light E -> electrical E to interpret as vision
== opsin prot + small mol ‘retinal’ (~vit A)
why does vit A deficiency cause night blindness
vit A has be converted -> retinal to make rhodopsin (all animals can do)
* night vision = most sensitive = lost 1st
* dogs/apes/horses can convert β-carotene -> retinol but cats can’t = need consume vit A in diet
photopigments in cones
diff opsins in diff colour cones as tuned pick up diff colours
1. ‘blue rhodopsin’ = opsin S + retinal
2. ‘green rhodopsin’ = opsin M + retinal
3. ‘red rhodopsin’ = opsin L + retinal
trichromats
= have 3 diff opsin containing cones - most primates, most other animals can’t see in 3 colours
other mammals inc ungulates, cats = 2 diff opsin-cont cones = dichromats
birds = tetrachromats as have UV asw
night vision
rods more sensitive light than cones so deer have more cones = better light sensitivity = better night vision
just diff adaptations: colour, night vision etc
photoreceptor signal transduction in darkness
high cyclic GMP (cGMP) = special Na+ channs open = Na+ thru -> cell = photoreceptor cells active = release glutamate (NT)
cellular anatomy photoreceptor
lamina of discs stacked to pick up as much light as poss w/in mem
* light detecting prot (e.g. rhodopsin in rods) + cGMP-gated Na+ chann embedded in disc mem together
* transducins (G prots) embedded in disc mem too
what happens when light falls on photopigment
- light E ‘trapped’ by cis-retinal => changes conformation -> trans-retinal
- no longer fits in large opsin = ‘falls out’ = opsin undergoes conform change = active
- active opsin causes α subunit of transducin to activate cGMP phosphodiesterase
- cleaves cGMP = decr cGMP conc (incr GMP) = Na+ channs close = cell hyperpolarises = no release glutamate
== photoreceptors deactivates by light == no glutamate
analagous to NT acting via G-prot receptor
role of glutamate in photoreceptor signal transduction
have ‘on’ + ‘off’ bipolar cells where glutamate inhibitory/excitatory so turned on/off by light
1. ‘on’ activates ganglion cell w light
2. ‘off’ activates ganglion cell in darkness
then axons of ganglion cells form optic nerve so having ‘on’ + ‘off’ means more info to brain to work out what image is