Biophysics of olfaction and taste

Biophysics of vision

Biophysics of Olfaction
Olfaction also known as olfactics; is the sense of smell. This sense is mediated by specialized sensory cells of the nasal cavity of vertebrates, and, by analogy, sensory cells of the antennae of invertebrates. Many vertebrates, including most mammals and reptiles, have two distinct olfactory systemsthe main olfactory system, and the accessory olfactory system (mainly used to detect pheromones). For air-breathing animals, the main olfactory system detects volatile chemicals, and the accessory olfactory system detects fluid-phase chemicals. For water-dwelling organisms, e.g., fish or crustaceans, the chemicals are present in the surrounding aqueous medium. Olfaction, along with taste, is a form of chemoreception. The chemicals themselves which activate the olfactory system, generally at very low concentrations, are called odorants.

The Olfactory Epithelium

In vertebrates smells are sensed by olfactory sensory cells (neurons) in the olfactory epithelium. The proportion of olfactory epithelium compared to respiratory epithelium (not innervated) gives an indication of the animal's olfactory sensitivity. Humans have about 10 cm2 (1.6 sq in) of olfactory epithelium, whereas some dogs have 170 cm2 (26 sq in). A dog's olfactory epithelium is also considerably more densely innervated, with a hundred times more receptors per square centimeter. Molecules of odorants dissolve in the mucus lining the superior portion of the nose cavity and are detected by olfactory receptor cells on the dendrites (olfactory sensory cells being kind of sensory neurons). This may occur by diffusion or by the binding of the odorant to odorant binding proteins.

Sensory transduction - general

Receptors transform an external signal into a membrane potential Two types of receptor cells: - a nerve cell - a specialized epithelial cell

Receptor adaptation - general

Tonic receptors
-- slow acting, -- no adaptation: continue to work for impulses as long as the stimulus is there (ex: proprioreceptors Proprioception is the sense of the relative position of neighbouring parts of the body )

Phasic receptors
-- quick acting, adapt: stop firing when stimuli are constant (ex: smell) with rapid variation of action potential to the beginning

Olfaction
Specialized neurons present in the olfactory epithelium in the nose. They project cilia into a mucus layer. The cilia are able to bind to odorant molecules the binding triggers an Action Potential which is transmitted to the olfactory area of the olfactory bulb olfactory cortex (lower frontal area and limbic system of the brain Each olfactory receptor is specialized for 1 odorant molecule

Taste receptor cells

Receptors for taste are modified epithelial cell present in taste buds located on the tongue, roof of the mouth and pharynx

Taste receptor cells for sweet and bitter ligands

 Cuplarea receptor-ligand duce la modificari locale ale campului electric dipolar al membranei Aceste modificari sunt capabile sa activeze enzime din membrana ca adenilat ciclaza (prin intermediul unei proteine de tip transducina-proteina tip G) care determina in citoplasma transformarea ATP (adenozin trifosfat sau acid adenozintrifosforic) in cAMP (adenozinmonofosfat ciclizat) Sub actiunea unei protein-kinaze, gruparile fosfat eliberate fosforileaza alt substrat proteic favorizand deschiderea unor canale de potasiu Ca urmare a modificarii locale a potentialului de membrana se deschid canale de calciu-sensibile la voltaj In final se elibereaza molecule de neurotransmitator in spatiul sinaptic= transmiterea informatiei

Principalele fluxuri de ioni care circula prin membrana celulara (R)

Transport pasiv=influx de sodiu, sau calciu precum si eflux de potasiu Transport activ = eflux de sodiu sau calciu sau influx de potasiu Din punct de vedere energetic: la tr. pasiv nu se consuma din energia metabolica dar la transport activ da (energia din moleculele macroergice de ex de ATPadenozin trifosfat) Transport pasiv prin canale ionice (agregate proteice) Transport activ prin pompe ionice (agregate proteice cu proprietati enzimatice)

Tendinta ionilor de sodiu (ca si de calciu) este de a intra in celule (transport pasiv prin canale ionice);

-transportul activ (prin pompe ionice) restabileste starea initiala deplasand ionii in sens invers gradientului electrochimic cu consum de energie din molecule macroergice cum sunt cele de ATP
Tendinta ionilor de potasiu este de a iesi din celule (prin transport pasiv)

Molecule macroergice
Moleculele macroergice din celulele organismelor vii au rolul de stocare a energiei provenit din procese metabolice de internalizare a energiei din exteriorul organismului (din degradarea alimentelor sau din fluxul energetic solar cum este cazul ATP la plantele autotrofe). Aceste molecule au i capacitatea de eliberare a energiei nmagazinate cnd procesele metabolice consumatoare de energie o cer. n lumea vie se cunosc mai multe tipuri de compui macroergici (tabelul *), care nmagazineaz energie chimic la nivelul unor legturi interatomice cum sunt: legturile hidrolizabile de tip P-O, care sunt i cel mai des ntlnite, (legturile P-O apar n cadrul gruprilor pirofosfat, acil-fosfat i fosfofenol); legturile hidrolizabile S-C, ntlnite mai rar, de exemplu n cadrul gruprii tioester, prezent la unul din metaboliii cei mai importani - acetil coenzima A. legturile hidrolizabile P-N, destul de rare, prezente n molecula de fosfagen (sau fosfocreatina).

Cel mai caracteristic compus macroergic este acidul adenozintrifosforic - ATP care prin hidroliza unei grupri fosfat trece n ADP - acid adenozindifosforic.
Tabel cu energiile degajate prin hidroliza unor compui cu fosfor (dup M. Isac, 1987)
Compus fosforilat DG(cal/mol) Compus fosforilat DG(cal/mol) Compus fosforilat DG(cal/mol) Compus fosforilat DG(cal/mol) Compus fosforilat DG(cal/mol) P-enolpiruvat 1,3-di-P-glicerat P-creatin Acetil-P ATP 7000 Fructozo-6-P 3800 3-P-glicerat 3100 12800 11800 10500 10100 Glucozo-1-P 5000 Glucozo-6-P 3300 Glicerol-1-P 2300

Molecula de ATP
Molecula de ATP prezint ncrcare electric (patru sarcini negative localizate pe legtura fosforic), toate cele patru grupri fosfat fiind disociate n condiiile pH-ului neutru (7,0) din citoplasma celular. Stabilitatea compusului macroergic ATP este dat i de interaciunile ntre dou grupri fosfat negative monovalente cu un cation bivalent de calciu sau magneziu. Din punct de vedere electrostatic msura energiei stocate n aceast molecul este dat de cea a energiei electrostatice a complexului de sarcini electrice negative, energie de respingere care este compensat la nivelul energiei interne a compusului stabilizat n care cele patru sarcini sunt meninute n apropiere una de alta.

Structura moleculei macroergice de ATP (acid adenozintrifosofric)

Reacia reversibil ATPADP joac rolul unei baterii energetice celulare care se ncarc prin fosforilarea ADP i se descarc prin hidroliza ATP:

ATP 4 H 2 O ADP 3 HPO 4 H

ATP este direct implicat n multe procese energetice din celul, cum sunt: glicoliza fosforilarea oxidativ fotofosforilarea fiind prezent n toate tipurile de celule animale i vegetale din scara filogenetic

 ATP se poate transforma n AMP ciclizat (cAMP) avnd rol n semnalizarea celular Guanozintrifosfatul (GTP) are un rol asemntor ATPului (de stocare de energie celular) iar cGMP este si el ntlnit n semnalizarea celular. Transportul activ transmembranar, contracia muscular i bioelectrogeneza sunt doar cteva din fenomenele biofizice n care astfel de molecule sunt, evident, indispensabile

Taste receptor cells for sour and salty ligands

 The mucus overlying the receptor epithelium contains mucopolysaccharides, salts, enzymes, and antibodies (these are highly important, as the olfactory neurons provide a direct passage for infection to pass to the brain). In insects smells are sensed by olfactory sensory neurons in the chemosensory sensilla, which are present in insect antenna, palps and tarsa, but also on other parts of the insect body. Odorants penetrate into the cuticle pores of chemosensory sensilla and get in contact with insect Odorant binding proteins (OBPs) or Chemosensory proteins (CSPs), before activating the sensory neurons.

Olfactory sensory cell (neuron) -with a single thick dendrite and an unbranched axon projecting to the olfactory bulb. The transduction region in grey - and signaling region in yellow. Transduction occurs in the cilia, which extend into the mucous layer. A receptor-coupled second-messenger system (next picture) results in the opening of a cation-selective channel in the ciliary membrane. The influx of cations depolarizes the cell membrane from its resting level near 65 to 45 mV, in a graded manner. This depolarization spreads by passive current flow through the dendrite to the soma. A depolarization that reaches 45 mV is sufficient to activate voltage-gated Na+ channels and initiate impulse generation. This Na+ current along with several varieties of voltagedependent K+ currents and a small Ca2+ current produce one or more action potentials that are propagated down the axon to the brain.

A receptor-coupled second-messenger system

G-transducina

 It is similar to the transduction mechanism for photoreceptors, in which the second messenger cGMP works by directly binding to ion channels, suggesting that maybe one of these receptors was evolutionarily adapted into the other. There are also considerable similarities in the immediate processing of stimuli by lateral inhibition. Averaged activity of the receptor neurons can be measured in several ways. In vertebrates responses to an odor can be measured by an electro-olfactogram or through calcium imaging of receptor neuron terminals in the olfactory bulb. In insects, one can perform electroantenogram or also calcium imaging within the olfactory bulb. The receptor neurons in the nose are particularly interesting because they are the only direct recipient of stimuli in all of the senses which are nerves. Senses like hearing, tasting, and, to some extent, touch use cilia or other indirect pressure to stimulate nerves, and sight uses the chemical rhodopsin to stimulate the brain.

Electro-olfactogram

Typical electro-olfactogram (EOG) responses elicited by the monoamine and polyamine odorants and the amino acid standard L-glutamine.

Michel W C et al. J Exp Biol 2003;206:1697-1706

Spermine is a polyamine involved in cellular metabolism found in all eukaryotic cells. Formed from spermidine, it is found in a wide variety of organisms and tissues and is an essential growth factor in some bacteria. It is found as a polycation at physiological pH. Spermine is associated with nucleic acids and is thought to stabilize helical structure, in particular, in viruse

Anosmia is a lack of functioning olfaction, or in other words, an inability to perceive odors.

(A) Representative electroolfactogram traces from WT (n = 9) and rd16 (n = 5) mice after exposure of the olfactory epithelium to three concentrations of amyl acetate -rd16 mice are anosmic.

Biophysics of Vision

Eye structure and sensitivity Optics of visual system in humans Retina structure fotoreceptor cells

Eye structure and sensitivity

 The eye can only perceive a small portion of the spectrum of electromagnetic waves

 In order to see an object: - 1- the pattern of the object must fall on the vision receptors (rods and cones in the retina) accommodation - 2- the amount of light entering the eye must be regulated (too much light will bleach out the signals) - 3- the energy from the waves of photons must be transduced into electrical signals - 4- The brain must receive and interpret the signals

Optics in human eye

It is the process of adjusting the shape of the lens so that the external image fall exactly on the retina

Accommodation
Object is far the lens flattens Object is near the lens rounds

Figure 10.25

Accommodation abnormalities
Myopia Hyperopia Astigmatism: the cornea is irregular irregular pattern of vision Presbyopia: stiffening of the lens occurring with aging increased difficulty with near vision

Figure 10.27ab

Figure 10.27c

Regulation (adjusting) of the amount of light entering the eye

The iris controls the amount of light entering the eye cavities The contraction of radial or circular smooth muscles located within the iris permit changes in the pupil diameter

Figure 10.28a

Eye abnormalities
Glaucoma Cataract

Figure 10.27b

Retina structure photoreceptor cells

Three cell layers:
-- outer layer: photoreceptors- rods and cones

-- middle layer: bipolar neurons -- inner layer: ganglion cells

Phototransduction
Photons hit the pigment of a photoreceptor enzymes are activated in the cell which modify its state of polarization the signals are sent to visual area of the occipital lobe of the brain through the optic nerve

Figure 10.31

Figure 10.33b

Neural processing
The bipolar neurons and ganglion cells process the signal In the fovea where the acuity is the highest: 1 cone 1 bipolar cell 1 ganglion cell At the periphery: many rods 1 bipolar cell acuity is much decreased Other cells in the retina participate in signal processing

Neural pathway
The right visual field maps on the left visual cortex and vice versa

Figure 10.36

Neural pathway

What will happen if the left optic nerve is severed? What will happen if a person has a tumor in the pituitary gland (just below the optic chiasmata) and the inner fibers are destroyed?

What will happen if a person suffers a brain tumor on the right side of the brain around the lateral geniculate body?