TEMA 15: FOTOPERIODISMO, MELATONINA Y PINEAL.

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TEMA 15:

FOTOPERIODISMO,

MELATONINA Y PINEAL.

1.- Papel de la pineal y la melatonina en el sistema circadiano.

2.- Anatomía comparada de la pineal.

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1. PAPEL DE LA PINEAL EN EL SISTEMA CIRCADIANO

MAMIFEROS

VERTEBRADOS NO MAMIFEROS

“Zeitgeber interno”

Oscilador circadiano

(3)

Passer domesticus

PINEALECTOMIA Y TRANSPLANTE DE PINEAL (AVES)

Lesión

Transferencia de

periodo y fase !

DONANTE

RECEPTOR

(4)

Anfibios

Teleósteos

Lampreas

(5)

Cartílago

Vesícula

pineal

Habénula

Techo

óptico

Pineal de la lubina

(6)

Reptiles Aves

(7)

Nat

NA

MEL

Luz

Luz

RETINA

Nat

MEL

SCN

GCS AMPc

Verdaderos

fotorreceptores

Fotorreceptores

modificados

Pinealocitos

en sentido estricto

RETINA

SCN

GCS

Luz

Nat

MEL

AMPc Hiperpolarización

TIPOS DE PINEALOCITOS

MAMIFEROS

AVES

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SEROTONINA

SEROTONINA

MELATONINA

MELATONINA

(5-Hidroxitriptamina)

(5-Metoxi-N-acetiltriptamina)

NAT

NAT

(N-acetiltransferasa)

HIOMT

HIOMT

(Hidroxiindol-O-metil

transferasa)

N-ACETILSEROTONINA

N-ACETILSEROTONINA

(5-Hidroxi-N-acetiltriptamina)

RUTA DE SÍNTESIS DE MELATONINA

Hora del día

(9)

Daily (T=24h)

Tidal (T=12.4h)

Seasonal (T=365d)

Lunar (T=29.5d)

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Skin Craneal bone Midbrain PINEAL ORGAN Pinea tract Parapineal Dorsal sack

Phototransduction

Phototransduction

at the fish pineal

at the fish pineal

(H.W. Korf, Evolution of melatonin-producing pinealocytes, Plenum Press, 2000)

PINEALOCYTES

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Time of day

Time of year

d

d

DURATION

a

a

d

AMPLITUDE

Melatonin and phototransduction

Moon phase

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14:00 17:00 20:10 23:20 2:25 5:15 8:20 11:20 0,00 100,00 200,00 300,00 :5 :00 a a a a a c bc c a a a b b bc bc c Summer solstice a b d a b ac ac c a bd bd b ac 14 17 20 23 2 5 8 11 Time (hour) M el at o n in ( p g/m l)300 200 0 100

SOLSTICIO VERANO

SOLSTICIO INVIERNO

0 12 24

Hora del dia

15L:9D / 9L:15D

25.0

o

C

12.9

o

C

SÍNTESIS DE MELATONINA EN CONDICIONES NATURALES

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Environmental transduction

Environmental transduction

: daily,

: daily,

lunar and

lunar and

seasonal

seasonal

melatonin rhythms

melatonin rhythms

T e m p er a tu r e ºC 0 50 100 150 200 250 300 M e la to n in a ( p g /m l) JAN 13.2ºC FEB 11.5ºC MAR 13.6ºC APR 16.7ºC MAY 20.1ºC JUN 20.5ºC AUG 25ºC SEP 19.5ºC NOV 17.2ºC DEC 14.5ºC 0 5 10 15 20 25 a a a a a a a a a b b b b bc cd bc d bcd bc bc JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC

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Environmental transduction

Environmental transduction

: daily,

: daily,

lunar and

lunar and

seasonal

seasonal

melatonin rhythms

melatonin rhythms

JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC y = 8,2941x - 16,963 R2 = 0,8596 0 50 100 150 200 250 0 5 10 15 20 25 30 Temperature (º C) M e la to n in ( p g /m l)

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Environmental transduction

Environmental transduction

: daily,

: daily,

lunar and

lunar and

seasonal

seasonal

melatonin rhythms

melatonin rhythms

0 50 100 150 200 250 300 m e la to n in ( p g /m l) 13.2ºC 17ºC WINTER 16.7ºC 19ºC SPRING 25ºC 20ºC SUMMER 17.2ºC 18.4ºC AUTUMN a ab bc c a a b b a a c b

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Environmental transduction

Environmental transduction

: daily,

: daily,

lunar

lunar

and seasonal melatonin rhythms

and seasonal melatonin rhythms

0 50 100 150 200 250 MD ML MD ML MD ML MD ML M e la to n in ( µ g /m l)

Luz Lunar Natural Luz Lunar Artificial

b c a a a a a a 0 50 100 150 200 250 MD ML MD ML MD ML MD ML M e la to n in ( µ g /m l)

Luz Lunar Natural Luz Lunar Artificial

b c a a a a a a

Full moon

0 20 40 60 80 100 120 140 160 MD ML MD ML M e la to n in ( p g /m l) a a a b 0 20 40 60 80 100 120 140 160 MD ML MD ML M e la to n in ( p g /m l) a a a b

New moon

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Reproductores

Reproductores

lunares

lunares

Siganus guttatus (1

st

WCC, 2003)

Sugama et al., J. Pineal Res. 45:133-141, 2008

(18)

Melatonin binding sites

Melatonin binding sites

:

:

where does melatonin act?

where does melatonin act?

Oliveira et al., Chronobiol. Int. 25:645-652, 2008

(19)

Environment

(Light/temperature)

eyes

Neuronal Melatonin

Hypothalamus

(GnRH)

Pituitary

(gonadotropins)

PINEAL

+

-GÓNAD

Sexual Steroids

Sexual Maturation

Pineal

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LUGARES DE PRODUCCIÓN DE

MELATONINA

• PINEAL

– Secretada a la circulación y al LCR

• RETINA

– Acción local, adaptación a la luz y oscuridad,

protección radicales libres? Depende de la especie

• TRACTO GASTROINTESTINAL

– Acción protectora local y sincronización de ritmos a

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Carassius auratus

RITMOS DIARIOS DE SECRECIÓN DE MELATONINA

Dicentrarchus labrax

Sánchez-Vázquez et al., J. Comp. Physiol. 167:409-415, 1997 Iigo et al., J. Biol. Rhythms

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PLASMA

M

E

L

A

T

O

N

IN

(

p

g

/m

l)

MD +LIGHT

0

60

120

RETINA

MD +LIGHT

M

E

L

A

T

O

N

IN

(

p

g

/o

jo

)

0

1000

2000

3000

Influencia de un pulso de luz brillante en MD

(23)

Pineal

Pineal

in vitro

in vitro

culture

culture

:

:

melatonin production persist?

(24)

Lamprea japonica

CULTIVO IN VITRO

DE PINEAL

LD

Cultivo

estático

(25)

Solea senegalensis

0 50 100 150 200 27 39 51 63 75 87 99 111 123 135 147 159 171 Time (h) M e la to n in ( p g /m l/ 6 h )

(26)

RITMOS CIRCADIANOS DE MELATONINA IN VITRO

LL

Esox lucius

DD

Bolliet et al., J. Pineal Res. 16:77-84, 1994

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(Porter et al., J. Fish Biol. 58: 431-438, 2001)

Salmón atlántico

Influencia de la INTENSIDAD luminosa

sobre la síntesis de melatonina

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PLASMA

ML MD 0.6 2.4 60 600 50 100 150 200 250 300 350 400 M el a to n in (p g /m l) (7) (6) (8) (7) (8) (8) (7) a a a a b b b 6

Influencia de la INTENSIDAD luminosa

sobre la síntesis de melatonina

Lubina

Umbral = 3 Lux

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6 18 6 18 Time of day 6 18 6 18 3 0 0 : 0 lx 3 0 : 0 lx 3 : 0 l x 0. 3: 0 lx 0: 0. 3 lx 0 50 100 150 200 250 300 350 M E L A T O N IN L E V E L S ( p g /m l) PLASMA

ML MD PULSE PULSE PULSE PULSE 0,3 lx 1 lx 3 lx 1400lx

Umbral = 0.3 Lux

Tenca

Influencia de la INTENSIDAD luminosa

sobre la síntesis de melatonina

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Lenguado

Influencia de la INTENSIDAD luminosa

sobre la síntesis de melatonina

0 50 100 150 200 250 300 350 400 450 ML MD 3.3 5.3 10.3 51.9 P la s m a m e la to n in ( p g /m l) a a a a ab b Light intensity (µW/cm2) 2100 0300 0900 1500 1800 0000 0600 1200 P la sm a m e la to n in (p g /m l) 0 20 40 60 80 100 120 Time (h) a c ab bc bc a a

Bayarri et al., Physiol. Behav. 81:577-583, 2004

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Espectro solar que llega a la Tierra

Espectro solar que llega a la Tierra

E n e rg ía lu m n íc a (m ic ro W /c m 2 ) Longitud de onda (nm)

Absorción del

Espectro

Absorción del

Espectro

400 500 600 700 0 15 30 45 60 75 400 500 600 700 25 0

P

ro

fu

n

d

id

a

d

(

m

)

Longitud de onda (nm)

AGUAS OCEANICAS AGUAS LITORALES Y CONTINENTALES

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ML MD R 2.4 R 6 G 2.4 G 6 B 2.4 B 6 0 50 100 150 200 250 300 M el a to n in (p g /m l) b ac c c c ac b a (7) (8) (8) (8) (6) (7) (8) (8)

PLASMA

Lubina

0.0 1.0 2.0 3.0 390 426 462 497 533 569 605 641 677 713 748 Wave length (nm) Ir ra d ia n c e ( % ) Daylight Blue Green Red Bayarri et al., J. Pineal Res. 32:34-40, 2002

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0 50 100 150 200 250 300 350 400 450 ML MD 0,3 1 3 P la s m a ti c m e la to n in ( p g /m l) a a a ab b

Lenguado

0.0 1.0 2.0 3.0 390 426 462 497 533 569 605 641 677 713 748 Wave length (nm) Ir ra d ia n c e (% ) Violet Red 355 369 600

red vio white

a a

ab

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REFERENCIAS

 García-Allegue,R., Madrid,J.A., and Sánchez-Vázquez,F.J., 2001. Melatonin rhythms in European sea bass

plasma and eye: influence of seasonal photoperiod and water temperature. J. Pineal Res. 31, 68-75.

 Iigo,M., Furukawa,K., Hattori,A., Ohtani-Kaneko,R., Hara,M., Suzuki,T., Tabata,M., and Aida,K., 1997. Ocular

melatonin rhythms in the Goldfish, Carassius auratus. Journal of Biological Rhythms 12, 182-192.

 Iigo,M., Sánchez-Vázquez,F.J., Madrid,J.A., Zamora,S., and Tabata,M., 1997. Unusual responses to light and

darkness of ocular melatonin in European sea bass. Neuroreport 8, 1631-1635.

 Korf, H.W., 2000. Evolution of melatonin-producing pinealocytes, in: J. Olcese (Ed.), Melatonin after Four

Decades, Kluwer Academic/Plenum Press, New York, pp. 17–29.

 Oliveira,C., López-Olmeda,J.F., Delgado,M.J., Alonso-Gómez,A.L., and Sánchez-Vázquez,F.J., 2008. Melatonin

binding sites in Senegal sole: day/night changes in density and location in different regions of the brain. Chronobiology International 25, 645-652.

 Oliveira,C., García,E.M., López-Olmeda,J.F., and Sánchez-Vázquez,F.J., 2009. Daily and circadian melatonin

release in vitro by the pineal organ of two nocturnal teleost species: Senegal sole ( Solea senegalensis) and tench (Tinca tinca). Comparative Biochemistry and Physiology A 153, 297-302.

 Oliveira, M. T. Dinis, F. Soares, E. Cabrita, P. Pousão-Ferreira, F. J. Sánchez-Vázquez, 2009. Lunar and daily

spawning rhythms of Senegal sole Solea senegalensis. Journal of Fish Biology 75, 61-74.

 Samejima,M., Tamotsu,S., Uchida,K., Moriguchi,Y., and Morita,Y., 1997. Melatonin excretion rhythms in the

cultured pineal organ of the lamprey, Lampetra japonica. Biological Signals 6, 241-246.

 Sánchez-Vázquez,F.J., Iigo,M., Madrid,J.A., Zamora,S., and Tabata,M., 1997. Daily cycles in plasma and ocular

melatonin in demand-fed sea bass, Dicentrarchus labrax L. Journal of Comparative Physiology B 167, 409-415.

 Sugama, N; Park, JG; Park, YJ, Takeuchi Y, Kim SJ, Takemura A, 2008. Moonlight affects nocturnal Period2

transcript levels in the pineal gland of the reef fish Siganus guttatus. Journal of Pineal Research 45, 133-141.

 Vera,L.M., De Oliveira,C., López-Olmeda,J.F., Ramos,J., Mañanós,E., Madrid,J.A., and Sánchez-Vázquez,F.J.,

2007. Seasonal and daily plasma melatonin rhythms and reproduction in Senegal sole kept under natural photoperiod and natural or controlled water temperature. J. Pineal Res. 43, 50-55.

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REFERENCIAS

 Bolliet,V., Bégay,V., Ravault,J.P., Ali,M.A., Collin,J.P., and Falcón,J., 1994. Multiple circadian oscillators in the

photosensitive pike pineal gland: A study using organ and cell culture. J. Pineal Res. 16, 77-84.

 Porter,M., N. Duncan, S. O. Handeland, S. O. Stafansson, N. R. Bromage, 2001. Temperature, light intensity and

plasma melatonin levels in juvenile Atlantic salmon. Journal of Fish Biology 58, 431-438.

 Bayarri,M.J., Madrid,J.A., and Sánchez-Vázquez,F.J., 2002. Influence of light intensity, spectrum and orientation

on sea bass plasma and ocular melatonin. J. Pineal Res. 32, 34-40.

 Bayarri,M.J., Muñoz-Cueto,J.A., López-Olmeda,J.F., Vera,L.M., Rol de Lama,M.A., Madrid,J.A., and

Sánchez-Vázquez,F.J., 2004. Daily locomotor activity and melatonin rhythms in Senegal sole (Solea senegalensis). Physiology and Behavior 81, 577-583.

 Herrero,M.J., Madrid,J.A., and Sánchez-Vázquez,F.J., 2003. Entrainment to light of circadian activity rhythms in

tench (Tinca tinca). Chronobiology International 20, 1001-1017.

 Oliveira,C., Ortega,A., López-Olmeda,J.F., Vera,L.M., and Sánchez-Vázquez,F.J., 2007. Influence of constant

light and darkness, light intensity, and light spectrum on plasma melatonin rhythms in Senegal sole. Chronobiology International 24, 615-627.

 Vera,L.M., López-Olmeda,J.F., Bayarri,M.J., Madrid,J.A., and Sánchez-Vázquez,F.J., 2005. Influence of light

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