Distribution of Neuropeptide Y Immunoreactivity in the Brain Stem and Spinal Cord of the Rat

y 品 v N(~{.:~ gel可 t_ : ~ ~

',f ,tj

I)

i

~.;;

t r ilHJ t i 011 0

i N

e

u

r 仆 peptide

.y"

Irnn1unoreactivitv in the

F~rain

Steal

al1d SlJina l o r d of the

I之 at

Chin立 Liang

Shen

DeO,-:lItrne 什 t of /J口;，) t ηIT! 句 ， College of Medicine

Nation，子 1 ['heη q Kunq Univε rSlt句

Iζ-..indn ， !diwan. f?epubl ic ηf China

Abstract ., UtI distc i. b 、 It ur 3 1 一、 ?、 『1 ， ltp p ? t l ry

(i'..T.PY) lik l:T1Elur::_-'lrcactlV1.r-V r-he

1-.,_, ，_三嗎?、

j i c.:.l ; r c'n '、 d spiηaI cord ('C the rat 、 arl

Ln~:-j'l\l:l\-~:isL (.[-)ρrTl 廿 1 I(lea 1 izιtiml of this p 佇 ptide

\，f;.，: 立已 。 6 r1 、 irm亡 d. U 片 J 門只 NPY ant i ~erurrl 、 Jt. IVa 只 pi'.'-"(、 iblc trJ [(1 ('21 工 zc til 亡 im :l;unc)reactivity in T11:l ι\l 仆 "j 、 l. 手 11 川、 di 亡耳 toe 司 ted ('xc!u'; 、 V 亡 1v i Cj t h f' ~， sf e In向 I'I-i is 仁、 1 ud ♂(j the Locu 丹 c ertllt?tl 討司 t h (. (f 匕 i 戶口 、 nc 1:~te τd j rc?t 1 仁 u 1.ar ·'lUC 1t_~U 泊， 且几 d l he 、 Iitτy :1 C 1eu s~ ν...rhi.e' h (.1f 亡 :h f dr- r::. 抗日 \~.'h 尸 t-t:' rn(I 廿 t ( dt γcnc> ;ιJ!1 、 lrlρτ 't:'，:.C' i'-~e runs 3rf.' Lee 市 tcd,. r [1

ddi， t J. c~ 甘、 Spl 口斗! 、 H1C] 尸 tIS 'and tract of thi晶

t r:. 芷江ρ1 九 31 ηerve 、 仁 he m].dl ine raphe nul'lei 正mei th 佇

汁~)rs 弓，'inI'n c! the ::;pi:--1al cord also cont 廿 [η ♂ d

!,

py

neu γ Ii 月 t::nml1 stdiηeei f beIs weγf' fσuη 可 Sfs\/eT'a!

于佇立"后 1~1 ← h() brβi 了 stem 刁 1;'0 spinal cord. T 于:e

r，'，仁、耳 t der 只 rJ."\1 NPY-J. ik.e 'fib ♂「另 IV在了佇 10(' 迂 tedin the I! ;tιI.λf 、、 ~ 月 ~J 1 汀八斗 九 ue' tcu::3 a 口 d t r 品 t- t.he

(γ ] t.:(:.0 f lO;~-l 一←一 γH_, :-(iγfit 、!、 V

"、‘ ι、 i 。E'c]侶 。 f 九己只 us 斗 and

r

66 I

t:

;出一!迋

Some Giiiψr'bl， di 也 rJ' i but 工可 area 污 bc:t\i\/eeil !)("

catecho] ami nergi c andNPY t i be rs \,It' re d;:-;,:usscd"

The [unctio 口 s of the NPY neurons were also bri 佇 I J 、

discussed.

INTRODUCTION

Neuropεptide Y (NPY) was tiona] siιnifi" I t' Ciln then bθ

1 日 01 在 tr于 d from the pro 正 In 佇 braln ev.::~luιit 泣，

by usin倍 '~ chemi Cd! 志只月 ay r{.~r

之 l

C-terminal amid佇且 , 只 ad-r 1m MATEHIALS AND M 此 T 付。 DS

rnunoass 在 y 只r. udie 只 I; 什 Vγ du 1 t ‘!!!)i「 t 仆 (dr-", ".Ie1.-C'

d 佇 monstrated that NPY eli"l r t 正 d t r:3l'.Jγdial' PI ::-tu:-~~iun

butes i 口 the m 尸 mr:'aLian ral l·JLth peritldatρ/1v~; in 對 Iparafor一

口 ervous system In 亡 Ludi ，;立 brain 111廿 ldehvde (PL?) fixλt] Vl'. The

s t

fIHi ， 2 、為

Immunohistochemical

brai" S[(-"11I and

spi 川 a]

cor'd

(C:斗，

m

-1 ) λ 斗 AU e l 、 p frt. 、 ← f 』 fι 、 ρL nh 、i LEK -va c uw ), J 一 門\仆 ‘ y 7 」 i

,

QU 叮IA ft d LL VJ O I Da m e

-[ e e hu e v 社 1. S OL U HUA etL I YA LH C ιL TL

lnve 月 LIgate the NPY like seri 品 1iv with a ! reeχlng

FL T口已 心 U I F i l ιυ li f

e-'

QJ ' 吋毛 J ' 吋/」 '

yli

AYLn .、 4.1L va

--r

t'D C an ea

r--ol na um

--nunu ma 1 4 向“

rnic rotom、 jII t !I 廿 II 甘 [I/O 日 tal

plan 亡 Ti 出 5\H" 切扎討 incubated at e d , 1 • L ] I 、J 毛 、.J l ) I t y 缸 ，、 i 1 、J I l t -、

AL--ZT /斗

v

l l t 一 • L O

-P) 7 '、 UW Hd h

'

r e v e w 、 J FiL LU ' s e .可 i AU U +L QU

explored the NPY Lik 巳 IIWIIU 門 \"_;~ t 1 片 (~rum. r1 ituιHeel at I: 之 CJ()()

activity in the br 司 111 日rem. '['cle wi til a 門 :~:(\d l.. \ln! ‘ )~:pllatc'

ind 工 rect pcroxid品 se-'-anti De ['ox i htlCfer 、、 intdlnlll 已 n.) Trixtul1

dase method has been 正'mpL''-Vied rοX 一 lilO all (j ! U lH)γrn 、， I 日 od t

investigatθth亡 NPY i l1EllU1J 仆 sιtum ， 1'11 ! 1 恥、 :ue \..J;-~，(; 心 ('Ilh 扎 l 佇 d

reactivIty

. ! 9

lorelJrall1

1n rh (' "、 rpC (j-](){J) ~-j[·ld [)γ1 (IXLdase 一

Her 亡， d detail ",J d t. JP(_·t·{-:xlcla 丸"、 I: (/I))、 Of ， lpI ('xc S

analysi 只 of Lhe NPYi,!lJT11UnO roca(• f ()、 iO 的 1 ~1~立1'1<1 →[]j Ill.

Live ma.terials in the brain stc 口 r l 心 t)<o" t 1V 正 1 v~ 1"1d r 11L\r\ t)J:1(心 d ]11

and spi 口 a] co rd 0 f the r 社 L l:-i 1(:(J inJ. 仆 iιj saLured

i 0 日 1 、 A I rsk 、片 ι1 、 i 11 rt

-re t

-什d i f

--、 j

e-a AU fk x ι ﹒、 戶 r荒 、 、 l l 少' QU l 1iL 、 1 pi i 」 ι 、 • fL I I fAγ nh t vl ny

of the NPY--like peptid

,'

111 the U.! 九1 pfl()Sp}IdLv hufl-(~rι 八 fte r

亡 。 ntrλ L nervous s\lsLeT11 of Lh 廿 I() mInute估， J]I) 1][ or 30克

1伸串借此做;Y 在大白鼠腦幹和脊髓的分佈

The diaminobenz idine react ion I. NPY perikarya Ioeal i zation.

was monitored visually for 20 A) Midbrain:

67

minutes. Following the chromo一 In the midbrain

,

the

ge 口 reactio日， the tissue was labeled NPY neurons were found

throughly washed in O. I M phos- in the periaqueductal gray.

phate buffer

,

gelatin-coated

mounted on

slides

,

dehy-These NPY neurons did not appear in

either the whole rostrocaudal

drated in ethanol

,

cleared in

xylene

,

coverslipped with

extent of the central gray or in

the whole area in cross section

permount

,

and examined under the of the periaqueductal gray.

mlcroscope.

RESULTS

They were found in the caudal

part of the structure

,

mainly in

the ventrolateral part of the

In each brain stem and cerebral aqueduct (Fig. 2). A

spinal cord studied

,

NPY few cells could be seen in the

lmmu口 oreactlve perikarya and lateral ce 口 tra 1 gray

,

but none

fibers were prese口 t in the brain were observed in the dorsal part

stem and all levels of the of the periaqueductal gray.

spinal cord. The immunoreactive Several scattered NPY neurons

neuro口 s were mostly concentrated were occasionally seen in the

in the locus coeruleus

,

the rost ral inferior colI iculus as

ve 口 trolateral region of the well (Fig. ]). These cells were

medulla and the dorsal midline distributed in all layers of the

structures. The NPY-like immu- col1iculus.

noreactive fibers were B) Pons:

abundant ly present in the soli 一 In the pons

,

the locus

tary tract and nucleus

,

the coerulus was the 0 日 ly structure

Spl 口 al nuc1 巴 us and the tract of that was stained by NPY

the trigeminal nerve and the antiserum. Numerous locus

apex of the dorsal hor口， and the coeruleus

,

neurons

central gray posterior to the positive (Fig. 3).

were NPY

These NPY

central canal of the spinal neurons distributed throughout

cord. the whole rostrocaudal extent

,

GH

250 輯朋

F 1g.i . The dj 耳 tribut 1 n of perikarγ社'.v ithi 吉、七←汁 ferior colliculus

whJγι1 ex 卜 i t! t ';mmU110r 亡 deLlγsf 丹 1(11 NPY i.'; JIll 先 tτated (:-1γrc 叫 heads.) .λi 只 a I、 fli(:)~~n i ("'.:1 ( i('> f

figure ].

2

thμSOU 廿 rc 1n

65um

F·~2. l~ 'l'h r'JP~'" imnlunoreac'cive perik'aτya (司 τr(刊vhe 'ld 月) are J uLdLcd ]口

tf01 于 h9

織一

他

有叫咱

Fig.3.

in卜、 , )t ，~址是[ ?毛 1 i ( ,C,'cr u~，(>l~ ， 'J 丸 'I ,'-jt j:'i '、 ,::!ttl' C

70 沈 J古良

inferomedial

par

t.

A few cells

obex

,

a miximum

co 口 ce 口 tration

of

with

NPY

immunoreactivity

were

NPY

cells

was

observed

in

the

also

found

i 口 the

subcoeruleus

nucleus (Fig. 6).

The number of

region.

C) Medulla oblongata:

NPY-containing neurons decreased

in its caudal division.

The NPY

neurons

also

extented

laterally

to the medial vestibular nucleus

The

medulla

oblongata

con-

(Fig.

4).

Some

NPY

immunoreac-tained

the

largest

population of

t ive neurons were

also

found

i 口

NPY

positive

neurons

in

the

the

lateroinferior

part

of

the

brain stem.

However

,

the

motor

nucleus

of

the

vagus

,

majority

of

the

NPY

neurons

which is

adjacent

to

the

appeared

in

the

lower

medulla

solitary tract

nucleus.

The

caudal

to

the

facial

nucleus

.

nuc leus

0

f

the

spinal

tract

0

f

These

NPY 口 eurons

were

mostly

the

trigeminal

nerve

is divided

locat己 d i 口 the

ventrolateral

,

into

three

subdivisions:

the

la 仁 eral

and

dorsomidline

struc-

pars oralis

,

pars

interpositus

,

tures

(Fig.

4

,

5

,

6).

A

and

pars

caudalis

.

The

NPY

considerable

number

of

medium-

neurons appeared from the middle

sized

NPY

posit ive

neurons

were

part

of

th巴 pars

interpositus

found

throughout

the

whole

down

to

the

whole

of the

pars

exterlt of

the

lateral

reticular

caudalis

(Fig.6).

These

NPY

nuclear

regiorl

,

and

extended

neurons

were

small

with

pseu-Laterally

to

reach

the

nucleus

dounipolar

morphology.

NPY

01

the

spinal

tract

of

the

neurons

in

the

middle

inter-trigeminal

nerve

,

medially

to

positus

part

occupied the most

adJace口 t

to

the

inferior

01

ivary

dorsal

and

most

ventral

parts

nuel 己 us

and

pyramid

,

and

infe一 of

the nucleus. Then

,

gradually

riorly

to

lie

UpOrl

the

basal

the NPY neurOrlS appeared at the

surface of

the medulla oblongata

whole medial edge of the spinal

(Fig.5

).

The

rostral

part

of

nucleus of the trigeminal nerve

仁 he

nucleus

of

the

solitary tract

at

the

level

of the obex (Fig.

did not contain NPY "eurons (Fig.

6).

The

caudal

spinal

nucleus

(IHi ) ，仁\，為 fJr' 扣 fi品)心 11: 71

Fig.4. Me ()~ in _{I'C: 仁、 r}ripn ;;ho'v./s he NPY-like n 亡'urons

\,,;}rv fl 手Ja(~ι-'1 t1!('、() c'‘ : dtl~l;-:? ， ~/~1 斗芯， dnd ,., are high 可、(一可，、 it"Ji~' l 呼: 、 JF ' ，。 1 >1 P Y DOSi t:1ve 1"e act i ve neur ;:月 、 ee ft -4 、 「 Y 1, r: 1!l tht: rnedial vestibular

、 111(‘leu:: 心 I ，、 市 P 1 -ik.令 n 甘心了。 τI，S (~r ρnt vertically in τaphe

ma}2.fJ I In (y 】 γ 立 f tlil" prt~'p( 古 ctus hypoglossal

1 、 Ul' j(';1~; 耐 1\'1A '、、內 1 f 叫 j_ d1. t(J信 C 1 亡 u] us

,

iυn=inferior

It.~ JT:

v

~

).

>'1' 、 11La~~γ 人 i、J 、、l 、 也U刮 、 ‘、 袒 戶 ~Tnu 、，土 1，'I ?島、 i 、 d 、 7~ ir ♂ 寸 、， ~L f\ 4.

-Y u ! ";亡。 h ← x. 芯 5 人(:1U\、 J d 月 f ..乏 )'.JCj e. 、 c·i.- 1;1" ζ 、可 、i CJ C1 、 j '=-以 I 〔品← C:: 1 _{)告 :--La; t} -.rac:L l t In p,Jgi S 咒~-~J 1.1 小 i 正 i ，:--， 昀砂v ;._~1 t,'J.rv u. !"i -.，_~V( 、

rh 「卜位人川、川 1 、

FA 戶

緝毒13甘、

韋拉

,.

、叭"戶可 、 4 猩紅 「位叫什心 Vv 叫Z 可→94 卡， 入戶品已 F ，~可 〉寺、-會 區 "扭的 Z 電學 4 φ p

s

一刀尤 叫〔 ih 卅 【戶，一〕可 一 F心J 〉品一: 斗 P，'‘叭、 dF 之 FK H r 弋}

.

' F 』、4 CJ 紗z; _f且 戶"" 門~ 1; .，~ 命 l _‘一， 、~ ~ '-J 斗J 力 .J _戶。 >--, J ;~J. F iJ'.; :,.>' 心二〉 、μ _'一

-J~ r.r:; 斗"- 二) : 仆地, f戶4 的 _心呵

-

" r: j、I 1吋 戶G CJ { ~ 、也r '\,.,5 M 丹、 、ν " > ',0 6戶一4 已- '、 戶戶 F呼 _~.J.{ 、、 OJ 爪 ，~ !凸 山 J 只 '0 •4 e' 、也 戶 t "、 戶 4 ~ _、 ,-'JJ 、--cJ: ,,1" y f已 " _、J ,> :毛 ~

.-u、 合吋 ‘,:: /) 叫 ] 3 ...., .. 1、J 叮 be c f已 \0 ~ "

-

,...l-, 川、ι …吋 ,- ~ :::...; 戶可 ?叫 一ι 、肉叫 p , ); ?正士 U' ," F “ _外I、" 門 J _ν一、 " 、h、‘也 工7_已~ 已鬥 恥 J _,. 一J 可唱J 」 明、』 _門 們以 hr f -- _'1J 叭, _c rf:. 0-(-.d)

74

it":J .1:這

7

'

,

7A~

Fig. 7. NPY immunoreactive fibers form a dense plexus in the 廿 pex 0 f

the dorsal horn and moderately in the central gray

posterlor to Uje ce 口 tra1 gray. A few NPY neuro 日 S

(arrowheads) are found in the (、 entτa1 ,gray posterior to the

central canal (Fig. 7A). Lot of NPY fibers make axosomatic

synapt i c patter口 s in the ve 口 trolateral motor neurons (Fig.

可司司

神經性且大 Y 在大白鼠腦幹和脊髓的分佈

neurons which were mostly

located at its lateral edge

between the nucleus and the

spinal tract of the trigeminal

nerve. These were small in

size. A few NPY neurons were

also found in the spinal tract

,

in its dorsal part

,

at the

level of the nucleus inter一

pos Hus.

75

cervical segments. They were

small neurons with a morphology

similar to those in the spi 口 a1

nucleus of the trigemi口 al nerve.

In the sacral cord

,

mediun~sized

NPY stained neuro口 s appearecj in

the posterior gray c 心 fDmlssurθ

(Fig. 7). No NPY immunorθ active

perikarya were found in the

posterior central gray i 口 ttl('

The raphe nuclei contained cervical

,

thoracic or lumbar

a few NPY neurons which were segments.

restricted to the area between

the level of the facial nucleus II. NPY immunoreacti νe fibers

and terminals:

and the level rostral to the

hypoglossal nucleus. The NPY

contalnl 口 g nuclei were the

nucleus raphe magnus

,

the

nucleus raphe pallidus

,

and th 亡

A) Midbrai月 and ρoη5:

In the midbrai口， the

central gray around the rostral

cerebral aqueduct had very

nucleus raphe obscurus (Fig.

4).

lightly labeled NPY fibers.

The NPY neurons were also fou日 d

1 口 the nucleus _{preposltus from the periventricular fiber}

These fibers were extensions

hypoglossi (Fig. 4); however

,

system. At the level of the

d e

--a [ nb

'

戶 L r u s cu

e-m m o c r 。

"-r e • L S o p e h 令 aL H .1i 可d G 、 aaa I u O ζ1 e r e vd ν I 1 H e , Lhu +』 石 i O e n O n

hypoglossal nucleus. The dorsal NPY fibers were found at the

column relay nucle 工 contained 口 o I 口 ferior edge of the commissure備

NPY neurons.

D) Spinal Cord:

A few fibers also appeared 廿 t

the superior edge of the

posterior commissure. Caudal to

I n t h e s pin a 1 cor d

,

the NP Y the po s t e rio r c0mm iss u re

,

NP Y

neurons were seen occasionally fibers decreased in number in

in laminae I

,

II and III of the the periaqueductal gray until

dorsal horn in ail levels of the the level of the caudal midbrain

it可 沈清泉

appeared agal 口 on _the

ventrolateral surtace _{。 f} the

perlaqLh~ductai gray. These NPY

fihers _{dis 正ippeared} In the

The innervation of NPY fibers to

the solitary tract 在 nd nue I eus

j ncreased in the mi dd 1C' and

白L LH 1 、 l/ /。 ' 「J ph 1IA hr /t 、、 a l 旬， i U AU e m

--a AU u a c

central gray of the pons. posterior commissural nucleus of

The rostral

thεsolitary nucleus and the

motor nucl eus of the vagus

medulla received very dense NPY fiber

β ) Medulla obloη gata:

。 blo 口 gata had fe\-i NPY lnnerva- inn巳 rvation. The postrema area

t ions. Few NPY fibers appeared also co 口 tained a light NPY fiber

i:l tbe centra 立 relY of thρupper distribution. In addit ion

,

the

1'lCdulla. pdr 仁 Lculariy _{Just spinal nucleus and the tract of}

Celllc! ιI to Lt-; JOl=US 仁。 εruleus. the trigeminal nerve also had a

F 仁\\' j t 打手 'i~: were found on the lot of NPY stained fibers (Fig.

6); but it did not distribute

C(jveυf fourth VEntricl巴，

a: 仁l 、L 內主 ti 、J heavier labeling

'-"''',IS fO Ll ηd In the preposltus

to the whole rostrocaudaJ

stru仁 ture. NPY fibers appeared

ilU \..' lψus of the hypoglossal nerve in the spinal nucleus and the

d'ld thιventromedial wall of the tract of the vagus nerve at the

central gray. Some fibers level of the obex where the

亡 xtended laterally to occupy the clos巳 d medulla began. NPY

area dorsal and r()stra] to the fibers appeared a 口 the medial

cnch 1e-::l 了一 n 日 cJc:us. Midline edge of the nucleus and the

行 truetures uf the medulla also adjacent area between the

hdd tiy labeled NPY fibers. nucleus a 鬥 d the tτact. 行 everal

了 hese stained NPY ~工 b Ee I'只 "vere labeled fibers and perikarya

~-; (:-n at t~. d 、 τsal ed 立(， cf the were also observed in the dorsal

py 了正lmid ， 社 nζ1 extended laterally part of the spinal tract (Fig.

i 仆 reach the medial edge of the 6). Then the labeled fibers and

pa 了 s raj is of the spinal cells gradually appeared

ilU 仁 Leus 仆 f the trigeminal nerve. laterally in the area between

八 t the I'廿日 tral medulla

,

a few the nucleus and the Lrac仁， and

NPY fib 七 rs vv' 亡 re alcoo observed in this pattern continued to the

fllr~ 最近此狀 Y If 大白血腦幹干n 特髓的分佈

ventral born (Fig.

n.

Few NPY

C)

Spinal Cord:

fibers were found to exte 口 d to

In the spi 口 al cord

,

NFl'

工 mmu口 oreactive fibers were seen

the IT!巳 di :cl l part of the antεrior

funicuLus, but no NPY fibers

at all levels of the spinal were found in the posterior or

cord. NPY fibeτs were concent- lateral columηof the spinal cord.

rated most heavily in the apex

(laminae I

,

II and III) of the

DISCUSSION

The present studv ha 只

dorsal horn and the area demonstrated that NPY i. mmunore 、 ie

adjacent to th 巴 central canaL

,

tive materials are labeled on the

espεcialLy the posterior gray perikarya and nerve fibers and

c omm iss u r e ( Fig. 7 ) . In the terminals. NPI' eon t3 in i[1 只

lumbar and sacral cords

,

small neurons are fou口 d in the Locus

arεas very densely labeled with coeruleus

,

tectum of the

NFl' fibers on the dorsal edge of midbrai口， ve 口 trolateral medulla

the central canal were observed ohLοngatι， 口 ucleu月 _。「 4

、

l i

b-(F i g.

7) .

Ma 口 y labeled NPY solitary tract

,

spLnal [τ1 呂 emi;1C--il

fibers were also found to extend nucleus and tr 司仁 t ， and spinal

from the posterlor gray cord. The J oeus coerulell 弓，

commlssure to reach the nucleus of the solitary tract

,

intermediolateral cell column iηand the rεglO口。 f the LHeral

the thoraci.e and J umbosacral ret icuI 品「九 ucleus of the medull 司

segme口 ts

,

par 仁 ieularly the have the h 工 ghest density of NFl'

latter (Fig. 7). A few NFl' immunoreactive cells. The NPY

stained fibers at the lumbar and Lmmu內心 sta 工 ned fib 已 rs are dens 亡 'v

sa 仁 ral 1 巳 vel x_fLe

-

-

ded 、3ell ahUe咀1i! e'α ，可_A 口 L•

-

-

e QU(、' 1in1+L技 rVJ 7A

-

ucl(、 et-4心J

e ) ftk s l 'C

antericrLy from the posterior and tract

,

the spinal nucleus and

c oIllmiss u raJ g ray t o t h e v e n t r a I traeιof the trigeminal 口 erv己，

and medial edge of the a 口 terior and the motor nucleus of lhe

horn. 1可 he NPY fibers made vagus.

axosomatlc sy 口 aptlc contacts The NPY-like i~nunoreac

with pregangL工 O 口 ic n euro IIs iηtivity in th 亡 mammalian brain

the intermediolateraJ nucleus stem has been studied 0ηlγi 口 the

23

78 沈 j持良

reported by Yamazoe 已 t

present study agrees with most were immunostained wi th NPY

antiserum. In the rat

,

the locus

S 令， L U 守J S7ι nτ'l ra

However

,

the NPY neurons

appeared In neither the

substantia nigra

,

the interpe一

coeruleus contained about 43% of

all 日 oradrenaline

t

neuro 日 s i 口 the brai 口 J

producing Neurons

dun 仁 ular 口 ucleus nor the pontine of the locus coeruleus have

structures except the locus elaborately branched axons which

co 巳 ruleus ， which contained NPY ramify within the whole central

neurons as reported by Yamazoe nervous system. Anteriorly

,

it

23

et al--. In the studies of the projects to the cerebral cortex

,

coexistence of the NPY and hippocampus

,

septum

,

basal

[ore-7

catecholamines

,

Everitt et al" brain

,

thalamus and hypothalamus

.

coeruleus

,

the

also reported that the locus Additionally

,

locus coeruleus

ventrolateral sends out descending fibers to

brain

region of the medulla and the the cerebe llum

,

solitary nucleus had a lot of

NPY neurons. The NPY-like

fib e r san d termi 口 als ， however

,

stern

sensory nuclei and spinal cord

II.18

, .

~ Interest ingly

,

the cere

bellum was not seen to exhibit

stem. The

are not reported as being any NPY-like immu日 oreactivity i 口

、 A I --ivyd afL rL-Ti 'DV -JL • L ec Ia te r 口 0 .ln u fL 訂 il rn 巴，­ QU ey rp pm-‘ _{distri 一}

but:ion in the spinal cord of explanation is that

tai 口 ing coeruleus neurons do not

seve1 在 I species including the

ral has been demonst rated by

Gibso口 p

t a ] 8 . T h e i r re sLI l t S

confj cmed the present study

,

that NPY一 l:ke fibers are

concentτated in the apex of the

the present or l‘ 2.9‘ 16 studies

",-,-,

In prevIous One reasonaLJle

NPY-con-project to the cerebellum whereas

the noradrenergic neurons do.

Surprisingly

,

Gibson et al8 found

that transection of the upper

spinal cord did not decrease the

ce 門 tr 司 l gray and also the

the posterior NPY immunoreactivity in the

spinal cord. These authors

dorsλ] hor 口，

para 日yrnpathetic preganglionic

neurons.

In the pt-eseηt study

,

many

neurons in the locus coeruleus

proposed that NPY fibers in the

spinal cord are of intraspinal

origin

,

instead of arising from

liJlii¥笠腔且大 Y 在大白鼠腦析和 1千髓的分有 ji ₇₉

containingηεurons in [he bτain motor nucleus of the vagus、 the

stern. Thus

,

the locus coerulear nucleus of the solitary tract and

projection to the spinal cord thεadjacent tegme口 turn ， including

may be maiηly noradrenergic the Iλteral tegme口 turn (A 1

,

A2

,

) 2

, )

5

The locus coeruleus projection aηd .43) The adrenergic

to the lateral thaIamie 口 ucleus neurons in the br 司 in are found

has recent1y been co 日 firmed by mainly in the lateral te 只 mental

Wheat-Germ aggl 叫 1rlln 叩門 jugated

to horseradish

peroxid吐出 e

i7. I n

(Cl) and

。 bIo口 gata (C2)

rostral rnedull a

of the caudal

the I ate r a L t hal am i c reg i0 日， medulla oblongata. Usiηg 1mmu口。一

there are nO NPY terminals. The histochemical identification of

presence of norepinephrine in tbe catecholami口 e synthesi只 lng

1 . e .

,

thyrosine

the sensory relay 口 ucJei of the enzymes

,

lateral thalamus has been de- hydroxylase

,

dopamine

monstrated by histofluoτescence -hydγoxylase and phenyl e thano] a 一

techn工 ques and by the immunohis

tochemica 1 ideηtification of the

J3

,

enzyme dopamin佇- -hydτoxyLase

20

,

22

There fore

,

locus

mine-N-methyi- transferase.

]0

Hokfelt 巴 t a1' ~ and Everitt et

a l7h ave d eIIIOHS t ra t e d t hFIt t TI e

NPY immunoreactive cell bodies

coeruleus neuro日 s coηtaining NPY are colocal ized in most noradre

seem not to project to the 口 ergic cell bodies of the AI/C]

1ater;:Jl thalamic l1tlc·jε 1.• The 只 roups 1n the ve 口 trolateral

Similarly

,

noradrenergic

,

rather than the medulla oblongata.

NPY

,

projection of the locus

NPY

immunoreactivity IS also

coeru leus to til ( ' 1 atera I gen i- ob 只 erved in the majority of the

culate 口 ucleus i 月 also co 門 fjrmed adrenergic perikarya of the C2

J J

, )

3 P I Trl

巾l _{distributio 口}

catecholamine-containiηg !l euro 汀 s coηtaining and

group. In the nucleus of the

of solitary tract

,

some 口 eurons

both NPY

in th 佇 ce 口 tra! nevous sv 污 tem has adreanal in have been found. the

4

, ]

4

,

l5

been well established"'" /\2 and C2 groups of somata in

戶、

-I

?i

noradrenergic neuronal catecholamine do not contain NPY

system origin技 tes m 丘 iη1y i n the neurons. The coexistence of both

~-~1 !-:< ,..;-,C 鼠。 口?

mod 'l lιti Ci1.1

;" t

l i: 仁

] T'i L 口 v (J !ved

;、'~~' peγ 、

rtIle:>tiS

、 c s !1C' l i

kl'

C(J;lduc 上 Jon 、 sCJr~' 丘iLJC 后 enc于 rιJ N' r:、 SUbCGl ru 1. u~~. the 門、 t fL -hu dorsal e LU Fi terrιD 已 raturε. pa 工 nand {了 ol L)--not do 廿 1 1(1 c βtccflolamines Le.S cord pJ i1 ιl t n• f、()1.UTI' 泣。! n、正支出 t之一('e PI:;il 日， LI]τ 1n l 只℃ c::1 nledu.l la 、 J bi III l1UC,LC' r 己 13 f[1reh 了在 1n~ 、! ) t i rk

-:i > l r 曰~-r(11 1 亢 par 只 th 。 hIe 日立在 t- i r a d i ()l. TnTT1U:i 仆 aS:3av us:n 名 t l et

r

什\l el C' llS S T) 1nell t I]C 。 r technique,';, 、L 、 lt I Pa i r )、 Jhyι ff '， l 勻， ι ( i rLd 鬼 門d m 7 ( γ4 1 and are, \.,rt-jl l.: 'fl nerve t γL 已正 i 川l.rJ λ1 the that feHl!l 、i l,' l 、!A Y 、 k il k、。、 A 。 CCS~至 urζi dnd touch i.r1 irJ\/U 1ved 亡 。 ntaL Cl J 口已 NPl' dεve I( 、 pme]lt h‘:lve noL do howev已 γ ， sensatl ns 、 the bral:l th 正 1[1 HKJ 1i !斗 、A t t lnn 世 rvatlon. NPY that to similar l S rat cmhryunie distrlhuLerj are fihe 了 S NPY catecholaminergic the 。 f tsi r l

-1 < !um 日 a l' 1c)wer the ln de 口 sely that found also They syst'_'I叮當 l 1i 仁 4t 、 intermediolateral sacral and st 己 m brain the 1!l NPY thf' 一、.、 :-')('drc'elv 。口lY and 亡。 lum日， 亡廿 rly very develop c!ie!l e' ρph'::llon 。 Ylldpt Lι t 「 l i i 仁 ord. thord 亡 1(' pc;. 品 tconception) ， i 斗 -da\P 、 . 、J ( . • ls f\ t (' rnl n aIs NP',! t t'i pat t' crη a play m 社 y NFY brA S 巴 rk fi ‘ 可 ug，試(:st ?-4 、

-7j a4 t、 the L ;,; 正 IIr\、T"lS ar 口 Et c! 、 J ) fL Y T、 '!、 AU t

-i et t \1 ;'ld clill(' n t :-1] also lS n 、1c1e 1.1 s 111tC 了 rn t'~(l .1d tεrλ! Dr-~:l n 吧 ((] t oh el a1S() m:"1y ! illers NP':{ !可干 內 iIi 屯， i 仆 ur~d ~ dIllon 心Jll.L、 the } n [nvo 1vet] L)t

!,'U NeTT ONAL CONS I DF:RATlON

L[1(0 'i n l)r 只{]1'1>~ ,0 (',f 1"f~ 只 U 1. dt i_on t'l':]\' 手4 rml l'、J 1 己 1'1(""C\i 14PY 1i i ( 1 T L 、l nH ) /、 1 、 )

av

-1 (-)"，~: t=、 r 社門 \1 pelvic 、 1[1 eL 凡 tribute 「、 3 h:之(一 "'V ( CJ '1 Dl GiS 亡。 vιr-c'(.1 J I ‘斗。 1 可 c av1.t J 巴", (',)rei 日 pInι! 1.Itt于 。i ιIpC'X til ( ( ;、 1 l •} Lt-)rnl1 汀 atι -I 1hers NPY 仁 [1at p l..rl:-j a 'Hi l y1 ) 1 1 dol-S 凸 A til ( 、: neur'OllS mot(、 γ V 已 1t r al l.amln~.:t e "亡 ~.1 S. t [' :';!(叮叮 、~1 、J­ t k f 1II 1 ) }

-4 IJ 11 -A r y--、 ( Iurnbosacral 出 p 1lai. l (' 'tll t,'V 1 ( f t fik tl 吉、)'[，.- 1(' 1γ Ct三 I" 'JIC:il nuc 1,(-jilS t r 1 PJ'ITl1n 凸[ c~;p:');~i 3i t 1 、巴 , 1 只 γndpt 1C aXOSO f1l、1t '1~, 正H1 c-·xhibil r 正 Lav S(~:-lsor\ Lγ[~t t l a i !]dlCdl(: wc.ule! Thi 討 Dat:tern. fr,)1!1 j np" L t 、台 1V(~ 、 A tf 、 '，、 1 e( 、叭 弋 、 , 、& 1 丸 '1\10]\led hu tiber 哼 fji1 t ;\lPY Lh 品亡 troni 社 it l-ent.s ~~( 11 汗、:、 pr:n1;"lr'v mulor :1 re 月 ulζi.L'i t r)E 1n 社 ηd head. 可 f:C 、 伊、 1 ,-i

-LTι 丸、 4 、 4 i 卡 iL rt lj l : (Jv'/(r 1 門口('rvat (~ the υl neurons \vhich st. r iatp muse les I 亡，在 V 1 丘之 -匙， η「 呵呵 J th 己~; l斗 that "cr、l ' tnd 、社 tp 、J t、 k 、 i t ] -vL T rUd" 。{_.口只 r nu 仁 i 丘]

神經 WH太 Y 在大白鼠腦幹和脊髓的分佈 ₈₁

distribution i 口 human brain.

Polak.

1983.

Neuropept ide Y

limbs. Due to the mass1ve

number of NPY 口 eurons 1n the

neostr1atum

,

Adrian et alI

suggested that NPY is of funda一

Nature

,

306: 584-586.

2. Allen

,

Y.S.

,

T.E. Adrian

,

T.J. Crow

,

S.R. Bloom

,

and

mental importance in the control

of motor funct ion. The

coexis-J.M. AIle口， K. Tatemoto

,

tence of NPY with the central J.M. Polak.

1983.

Neuropep一

catecholamine neurons

,

espe 一 tide Y distribution in the rat

cially in cell bodies containing brain. Science

,

221 :877-879.

adrenaline

,

suggests that NPY.

3.

Allen

,

Y.S.

,

T.E. Adrian

,

might function as a co-modulator

neuro日 S which are probably

1n some central adrenergic

J.M.. Allen

,

G.W. Roberts

,

T.J. Crow

,

K. Tatemoto

,

S.R.

Bloom

,

and J.M. Polak.

1983.

endogenous pp-like peptide

widely distributed i 口 the

involved in the regulation of

cardiovascular and respiratory

functions

,

and sleep and

wake-Neuropeptide-Y 1S the

fulness. NPY in the central brain of man and other

nervous system may have other mammals. J. Patho

l.,

140:129一

functional importance. These

130.

functio日 s of NPY remain to be

4.

Armstrong

,

D.M.

,

A.R. Chris一

clarified. topher

,

V.M.

Joh

,

and D.J.

Pickel

,

T.H.

Reis.

1982.

carried out in the laboratory of

DR. M.Y. Moore

,

Department of

Neurology

,

State University of

New York at Stony Brook. immunocytochemical

localiza-t ion of catecholamine biosyn一

Distribution of dopamine一，

noradrenaline-

,

and

adrenaline-contai口 ing cell bodies in the

Demonstrated the oblongata: by medulla rat was 1nvest 工 gat10n ACKNOWLEDGEMENT This REFERENCES:

I. Adria口， T.E.

,

J.M. Allen

,

S.

R. Bloom

,

M.A. Ghatei

,

M.N.

Rossor

,

G.W. Roberts

,

T.J.

Crow

,

K. Tatemoto

,

and J .M.

thetic enzyme. J. Compo

Neuro

l.,

212;

173-187.

5.

Dablstrom

,

A. and K. Fuxe.

1964 .

Evidence for the

82

conta 工 nlng 口 eurons 1 口 th 們

C 已 ntral nervous system I

Demonstration of monoamines

1n the cell bodies

brai 口 stem neuro 口 s. Acta.

Physiol. Scand 、 Suppl. ， 23:

2475一 2484.

6. Emso日， P.C. and M.E.

Quidt. 1984. NPY- a new

member of the pancreat1c

polypeptide family. Trends

Neuroscie口 ceo ]l: 31-35. 7. Everitt

,

B.J.

,

T. Hokfelt

,

-J I Tereηius ， K. Tatemoto

,

V. Mutt

,

and M. Goldstei 口，

]984. _{。 ifferential co-exis一}

沈清良

localization or Ileuropeptide Y

(NPY) i 口 the I'at hrain. SOC.

Neuroscie 口 C 亡， 9:291.

。 flO. Hokfelt

,

T.

,

T.M. Lundbeγ鼠，

K. Tatemoto

,

V. t-lutt

,

and L.

Goldstei. 1983. NPY and FMRF

amide neuropept ide--I jke

De immunoreactivities in catecho一

1amine neuro口 s of the rat

medulla oblongat正L Acta.

•

_{Physiol. Scand.}

_,

_117:315-318.

1I. Jone s

,

B . E . a 口 dR.Y

.,

Moore.

1977. Asce叫 1 月 projection of

the locus coeruleus in the

rat. An autoradio 只 raphic

(NPY) 一 like Immunoact1v1tvJ

tence of neuropeptide Y 12. Kalia

,

M.

,

D.J. Woodward

,

W.

study. Brai 口 Res. ， 127:23-53.

with catecholamines in the

central nervous system of the

τa t. Neurocience

,

II: 443一

462.

8. Gibson

,

S. J.

,

J.M. Polak 、

J.M. AlIe 口， T.E. Adria口，

J. 3. Ke II y

,

and S.R. Bloom.

1984. The distribut ion and

。 τIg1口 of a nove1 brain

peptid巴 ， nF opept ide Y

,

in

the 污 pin2 cord of several

mammals J . Compo Neurol.

,

227:78-:/1

K. Smith

,

and K. Fuxe. 1985.

Rat medulla oblo口 gata IV

topographical distribution of

catecholaminergic neurons with quantitative three-dimensional

computer reconstructIon. J.

Compo Neurol.

,

233: 350-364.

13. Lindvall

,

nu

A.

Nohin.

Bjoklu吋， a 叫 U. 只 tenevi.

1974. The adrener日 lC

lnnerva-t i0 口。 f the rat thaI amus <IS

r 亡 veλled by the glyoxyl ic acid

fluorescen仁 e method. J. Comp.

Neurol.

,

]54:317~3!， 8.

PoLak. and G.

,

Pelletier.

9. Guy

,

.f.、'l .S. ， Allen

,

1.3. 14. Moore

,

R.Y. and L.F. Kromer.

The organization of

1983

,

Immunocytochemical

197 日.

神經自生做 Y 在大白鼠Ii鼠幹和脊髓的分佈

system.

I 口 B.

Haber

and

vd 口。 h o o l s0 .YA 尸L ra 口 &m r A a 'H DA O H 盯 MLM

CEds.)

and

Behav 工 or.

Plenum

,

pp.77-87.

83

Distribution

of

neuropeptide

Y immunoreactivity

in

the

forebrain

of

the

rat

‘

Proc.

Nat

l.

Sci.

Counc.

B.

,

1]:

1]5一

15. Moore

,

R.Y.

and F.E.

Bloom.

127.

1979.

Central

catecholamine

20.

Swanson

,

L.W.

neuron

systems:

Anatomy

and

physiology

of

the

norepi口 e 一

pn rl ne

and

epInephrine

systems.

Ann.

Rev.

Neurosc ience

1

2:] 13- 168.

16. O'Donohue

,

Chranwall

,

T .L.

,

and

N.B.M.

D.A.

Dimaggio.

]983.

Distribution

of

neuropeptide

Y-like

immunoreactivity

in

the

rat

21.

Tatemoto

,

Complete

brain.

SOC.

Neuroscie 口 ce ，

9:

290.

17. Peschanski

,

M.

and

j .

Besson.

1984.

Coerulear

and

B.K.

Hartman.

1975.

The

ce 口 tral

-e , d 0 , nn nta u m-3 mf 叭叭 1 0 ，工 」 u n kao tyb .d mul et-L tse sc

y

sezi c) C 口 ( -le gcn rso 戶L 戶L-14 nrt eoa ruc dlo

af-their efferent connections in

the

rat

utilizing

dopami口 e

beta hydroxylure as a marker.

J.

Cowp.

Neuro

l.,

]63:467一

506.

Neuropeptide

K. Y:

1982.

amino acid

seque口 ce

of

the

brain

pept ide.

Proc.

Nat l.

Acad.

Sci.

USA.

,

79:5485-projections

to

the

lateral

5489.

diencephalon

in

the

rat.

An

22.

Ungerstedt

,

U.

anatomical

study

uSIng

wheat-germ

agglutinin

co 日 JU

gated

to

HRP.

Neurosci.

Lett.

,

46: 329-334.

197

J •

Stereotaxic

mapping

of

the

monoamine pathways in the rat

brain

,

Acta

Physiol.

SC2nd.

Suppl.

,

367: 1-48.

F.E.

] 974.

A

18. Pickel

,

V.M.

,

M.

Segal

,

and

23. Yamazoe

,

M.

,

S.

Shiosaka

,

Bloom.

radiographic

study

of

the

efferent

pathways

of

the

nucleus

locus

coeruleus

,

J.

Compo Neuro

l.,

155:15-42.

19.

Sh e

口， C. L. J

987 .

P.C.

Emson

,

and M.

,

Tohyama.

1985.

Distribution

of

neuro一

peptide

Y

in

the

lower

brai口 stem:

an

immunohisto 一

chemical analysi

日，

Brain Res.

,

84

沈鴻良

神經目生目太 Y 在大白鼠腦幹不日脊髓的分佈

;尤清良

中文摘要

本實驗以免疫組織化學法永研克大白鼠腦幹和脊髓內神經性 狀 Y 的分佈情形。利用 NPY 抗禮之研完發現 NPY 神經元廣泛的分 佈於腦幹中。它的分佈以豆豆斑枝，丹、網狀核區和孤獨核為最多 O 除夕|、三叉神經脊髓徑和枝，脊髓背角等也合并多的 NPY 神經元 O 至於 NPY 神經纖維亦分佈於腦幹和脊髓的并多區域 O 其中以 孤獨求，三叉神經脊髓徑和枝，迷走神經運動枝，脊髓背角頂部

為最多。本文討論了 Cat echolamine 纖維和 NPY纖維分佈土之差