墨龍與紅鯽的視網(wǎng)膜和視蓋解剖結(jié)構(gòu)比較

萬 安安樹青華田苗

(1. 南京大學(xué)生命科學(xué)學(xué)院, 南京 210093; 2. 安慶師范學(xué)院資源環(huán)境學(xué)院, 安慶 246000; 3. 安徽師范大學(xué)生命科學(xué)學(xué)院, 蕪湖 241000)

墨龍與紅鯽的視網(wǎng)膜和視蓋解剖結(jié)構(gòu)比較

萬 安1,2安樹青1華田苗3

(1. 南京大學(xué)生命科學(xué)學(xué)院, 南京 210093; 2. 安慶師范學(xué)院資源環(huán)境學(xué)院, 安慶 246000; 3. 安徽師范大學(xué)生命科學(xué)學(xué)院, 蕪湖 241000)

墨龍是一種由紅鯽進(jìn)化來的龍睛種金魚(Carassius auratus)。隨機(jī)取體長10—12 cm, 重約35 g的墨龍和紅鯽各4尾, 解剖取出整個(gè)眼球及腦, 并常規(guī)石蠟切片, HE染色。在光學(xué)顯微鏡下觀察墨龍和紅鯽的視網(wǎng)膜、視蓋系統(tǒng)的顯微結(jié)構(gòu)變化并比較各層厚度, 發(fā)現(xiàn)與紅鯽相比, 墨龍視網(wǎng)膜的總厚度下降29.9%, 其中外網(wǎng)狀層厚度增加2.5%、內(nèi)網(wǎng)狀層厚度增加11.8%; 而內(nèi)核層厚度下降21.6%、外核層厚度降低35.6%, 神經(jīng)節(jié)細(xì)胞層、桿錐層也變薄, 且后兩者分層不規(guī)則; 墨龍視蓋壁整體厚度增加28.9%, 其中除圍腦室層厚度減少22.6%外, 中央纖維層厚度增加12.8%, 中央細(xì)胞層厚度增加30.6%, 表面纖維層厚度增加21.9%, 且纖維遠(yuǎn)較紅鯽密集, 視神經(jīng)層厚度增加91.7%, 邊緣層厚度增加35.6%。結(jié)果表明長期的人工選擇不但改變了墨龍的外形, 而且使其中樞神經(jīng)組織結(jié)構(gòu)也發(fā)生了較大變化, 并推測(cè)墨龍的眼球直徑及視網(wǎng)膜面積較大, 從而導(dǎo)致自視網(wǎng)膜傳入視蓋的纖維增多, 是視網(wǎng)膜和視蓋中的傳遞神經(jīng)沖動(dòng)的神經(jīng)元、神經(jīng)纖維所在層段增厚的主要原因; 同時(shí)墨龍視網(wǎng)膜中色素上皮層向桿錐層交錯(cuò)對(duì)插, 富含神經(jīng)元的視網(wǎng)膜外核層、內(nèi)核層以及視蓋中的圍腦室層厚度也降低, 可以減少因視網(wǎng)膜面積大而造成的強(qiáng)光傷害; 此外由于墨龍的圍腦室層厚度降低, 導(dǎo)致其游動(dòng)及平衡能力較紅鯽差。

金魚; 墨龍; 紅鯽; 視網(wǎng)膜; 視蓋

金魚也稱“金鯽魚”, 屬鯉形目, 鯉科, 鯉亞科,鯽屬, 與鯽魚同用一個(gè)學(xué)名(Carassius auratus), 研究表明,銀灰色的野生鯽魚變?yōu)榧t黃色的紅鯽(Red crucian), 然后再經(jīng)過不同時(shí)期的馴養(yǎng)和選擇培養(yǎng),由紅鯽逐漸變成為不同品種金魚, 各種金魚的體態(tài)出現(xiàn)了與自然環(huán)境下的魚類明顯不同的特征[1]。除外貌特征的改變外, 有學(xué)者發(fā)現(xiàn)墨龍(Black dragoneye)眼球體積及視網(wǎng)膜整體面積較紅鯽大, 而厚度較薄[2]。鑒于此, 通過實(shí)驗(yàn)觀察, 本文對(duì)金魚中墨龍和紅鯽的視網(wǎng)膜和視蓋系統(tǒng)的顯微結(jié)構(gòu)進(jìn)行進(jìn)一步比較,以期有所發(fā)現(xiàn)。

本研究采用石蠟切片和HE染色技術(shù),通過光鏡對(duì)紅鯽和墨龍兩種金魚的視網(wǎng)膜和中腦視蓋的組織學(xué)結(jié)構(gòu)進(jìn)行了觀察, 為深入研究長期人工選擇條件下金魚的感覺器官和神經(jīng)系統(tǒng)的生理結(jié)構(gòu)變化提供基礎(chǔ)資料。

1 材料與方法

實(shí)驗(yàn)用金魚100尾, 購自安慶市花鳥魚蟲市場(chǎng),實(shí)驗(yàn)前置于實(shí)驗(yàn)室禁食馴養(yǎng)48h,隨機(jī)取體長10—12 cm, 重約35 g的各4尾置于4℃水中麻醉, 用游標(biāo)卡尺測(cè)量體長, 眼球直徑, 從第3、第4脊椎處剪斷, 將頭部轉(zhuǎn)移到Olympus SZ1145型體視顯微鏡下進(jìn)行解剖, 將整個(gè)眼球及腦取出后, 迅速用4%多聚甲醛的PBS溶液固定48h。用PBS清洗后, 按常規(guī)方法制作石蠟切片, 連續(xù)冠狀切, 厚度6 μm。

將所得切片做HE染色, Olympus BX51型顯微鏡觀察并拍照, 每張切片上隨機(jī)選取8個(gè)視野測(cè)量視網(wǎng)膜、視蓋總厚度及各層厚度(Unint: μm), 并計(jì)算平均值。所有數(shù)據(jù)采用平均值±標(biāo)準(zhǔn)差表示, 用t-test進(jìn)行各指標(biāo)的顯著性檢驗(yàn)。

2 結(jié)果

2.1 體貌特征

測(cè)量結(jié)果顯示, 實(shí)驗(yàn)所用紅鯽和墨龍的體長相似, 但墨龍的眼球直徑明顯大于紅鯽(表1)。

2.2 視網(wǎng)膜厚度變化

組織染色顯示紅鯽的視網(wǎng)膜7層結(jié)構(gòu)清晰易辨(圖版Ⅰ-1), 而墨龍神經(jīng)節(jié)細(xì)胞層(Ganglion cell layer, GCL), 桿錐層(Photoreceptor layer, PL)和色素上皮層(Pigment epithelium layer, PEL)各層界限不分明, 厚度不易分辨, 其中色素上皮層向桿錐層延伸并交錯(cuò)對(duì)插(圖版Ⅰ-2)。

選擇兩種魚的視網(wǎng)膜總厚度以及較清晰的4層:外核層(Outer nuclear layer, ONL)、外網(wǎng)狀層(Outer plexiform layer, OPL)、內(nèi)核層(Inner nuclear layer, INL)和內(nèi)網(wǎng)狀層(Inner plexiform layer, IPL)進(jìn)行統(tǒng)計(jì)比較。與紅鯽相比, 墨龍的視網(wǎng)膜出現(xiàn)一些明顯的變化(表2、表3): (1)視網(wǎng)膜總厚度平均下降29.9%,與紅鯽相比具有極顯著差異(P<0.001); (2)其中外核層厚度平均下降35.6%, 具有顯著差異(P<0.05); (3)外網(wǎng)狀層厚度平均增加2.5%; (4)內(nèi)核層厚度平均下降21.6%, 具有顯著差異(P<0.05); (5)內(nèi)網(wǎng)狀層厚度平均增加11.8%; (6)外網(wǎng)狀層和內(nèi)網(wǎng)狀層占視網(wǎng)膜總厚度的比例較紅鯽均出現(xiàn)顯著增加; (7)神經(jīng)節(jié)細(xì)胞層, 色素上皮層和桿錐層, 各層厚度不易分辨。

表1 紅鯽與墨龍?bào)w貌比較Tab. 1 Comparison of physical appearance of black dragoneye and red crucian (n=4)

表2 紅鯽、墨龍視網(wǎng)膜各層厚度比較Tab. 2 Comparison of the thicknesses of retina layers in black dragoneye and red crucian (μm)

表3 紅鯽、墨龍視網(wǎng)膜各層在總厚度比值(相對(duì)厚度)比較Tab. 3 A comparison between the relative thickness of retina layers in black dragoneye’s and red crucian’s (%)

2.3 視蓋各層厚度變化

組織染色顯示兩種金魚的視蓋結(jié)構(gòu)清晰易辨,依據(jù)林浩然等[3,4]分6層(圖版Ⅰ-3、4), 而與紅鯽相比,墨龍的視蓋出現(xiàn)一些明顯的變化(表4、表5): (1) 視蓋總厚度平均增加28.9%, 與紅鯽相比具有極顯著差異(P<0.001); (2) 圍腦室層(Stratum periventriculare, SPV)較紅鯽薄, 厚度減少22.6%, 與紅鯽相比具有極顯著差異(P<0.001); 圍腦室層占視蓋總厚度的百分比較紅鯽顯著降低; (3) 中央纖維層(Stratum album central, SAC) 較紅鯽厚, 厚度增加12.8%; (4) 中央細(xì)胞層(Stratum griseum central, SGC) 較紅鯽厚, 厚度增加30.6%, 與紅鯽相比具有顯著差異(P<0.05); (5) 表面纖維層(Stratum fibrosum et griseum superficiale, SFGS)較紅鯽厚, 厚度增加21.9%, 與紅鯽相比具有顯著差異(P<0.05); 且兩者結(jié)構(gòu)質(zhì)地明顯有差異, 紅鯽此層纖維密度較稀疏(圖版Ⅰ-3、4)。(6) 視神經(jīng)層(Stratum opticum, SO) 較紅鯽厚, 厚度增加91.7%, 與紅鯽相比具極顯著差異(P<0.001); 視神經(jīng)層占視蓋總厚度的百分比出現(xiàn)顯著增加。(7) 邊緣層(Stratum marginal, SM)較紅鯽厚, 厚度增加35.6%, 與紅鯽相比具有顯著差異(P<0.05)。

3 討論

墨龍全體漆黑, 身體短而肥, 尾鰭分叉大, 喜靜, 不善游動(dòng), 由紅鯽馴化而來[1], 屬龍睛類金魚,其眼球過分膨大而突出于眼眶之外, 眼徑明顯大于紅鯽。魚類眼進(jìn)化的特點(diǎn): 一般白天活動(dòng)靠視覺攝食的魚類, 有突出的眼, 其直徑大約相當(dāng)于頭長的l/5—1/6; 而黃昏或夜間活動(dòng)、主要用視覺捕食的魚,或至少部分靠視覺攝食的深海魚類, 眼徑可達(dá)頭長的1/3—1/2[5]。墨龍眼徑大, 推測(cè)其視網(wǎng)膜和視蓋也應(yīng)隨之而發(fā)達(dá)。

實(shí)驗(yàn)發(fā)現(xiàn), 與紅鯽相比, 墨龍的眼徑較紅鯽大,視網(wǎng)膜面積也較大, 其視網(wǎng)膜卻出現(xiàn)小眼徑魚類具有的特征[3,9—12]: (1)視網(wǎng)膜壁層總厚度較紅鯽平均下降29.9%; 桿錐層(PL)、視神經(jīng)節(jié)層(GCL)變薄,分層不規(guī)則; 感光細(xì)胞胞體所在的外核層(ONL)較薄, 推測(cè)其視網(wǎng)膜的感光能力較弱; (2)具有傳遞神經(jīng)沖動(dòng)作用的雙極細(xì)胞、水平細(xì)胞所在的內(nèi)核層(INL)[3], 含大量傳導(dǎo)纖維的外網(wǎng)狀層(OPL)、內(nèi)網(wǎng)狀層(IPL)較紅鯽均顯著增厚, 說明墨龍的視網(wǎng)膜整體輸出的神經(jīng)沖動(dòng)較紅鯽多。

表4 紅鯽與墨龍視蓋各層厚度比較Tab. 4 A comparison of the optic tectum layers in black dragoneye and red crucian (μm)

表5 紅鯽與墨龍視蓋各層與總厚度比值(相對(duì)厚度)比較Tab. 5 A comparison between the relative thickness of the optic tectum layers in black dragoneye and that red crucian (%)

可見與紅鯽相比, 墨龍視網(wǎng)膜上感光細(xì)胞的減少, 造成對(duì)外界光線的敏感度和顏色分辨能力下降,但墨龍的眼球直徑較大, 因此導(dǎo)致視網(wǎng)膜傳入視蓋的纖維增多, 并可能是導(dǎo)致其視網(wǎng)膜和視蓋中的傳遞神經(jīng)沖動(dòng)的神經(jīng)元、神經(jīng)纖維所在層段增厚的主要原因。

與紅鯽相比, 墨龍的視蓋壁整體增厚28.9%,除圍腦室層厚度減少22.6%外, 其余各層均明顯變厚: (1)表面纖維層(SFGS)含有接受傳入聯(lián)系的神經(jīng)元和神經(jīng)纖維、視神經(jīng)的前枝纖維,以及一些水平細(xì)胞和較大的錐體神經(jīng)細(xì)胞, 視神經(jīng)層(SO)含有來自中央和兩側(cè)視束的視神經(jīng)纖維[3—8], 觀察到表面纖維層(SFGS)的纖維分布遠(yuǎn)較紅鯽密集, 且這兩層均較紅鯽增厚, 其中負(fù)責(zé)傳入視覺神經(jīng)沖動(dòng)的視神經(jīng)層(SO)厚度增達(dá)91.7%, 在視蓋中相對(duì)厚度也較紅鯽增大, 表明自視網(wǎng)膜輸入視蓋的神經(jīng)纖維數(shù)量確實(shí)增多。(2)除含少數(shù)零星的水平細(xì)胞外, 主要由傳出神經(jīng)纖維組成的中央纖維層(SAC); 含有錐體神經(jīng)細(xì)胞的樹突和胞體位于中央細(xì)胞層(SGC)的上行神經(jīng)纖維的邊緣層(SM)[3—9]各層厚度均增加, 說明墨龍自視蓋中發(fā)出的上行及下行神經(jīng)纖維數(shù)量增加。(3)視蓋的中央細(xì)胞層(SGC)內(nèi)含有多極神經(jīng)元、大梭狀神經(jīng)元和梨形神經(jīng)元等多種類型的中間神經(jīng)元[3—8], 墨龍的此層厚度增加說明其視蓋各層間信息交換的能力增加。(4)墨龍圍腦室層(SPV)由圍腦室神經(jīng)元和視蓋深層纖維所組成, 神經(jīng)元發(fā)出的神經(jīng)纖維向外穿過視蓋各層, 廣泛地與視蓋各層發(fā)生聯(lián)系, 并可能是視覺信息傳出的主要途徑[3,7,9]。而墨龍的圍腦室層的實(shí)際厚度和在視蓋中的相對(duì)厚度較紅鯽均顯著減少, 表明其信息處理能力較紅鯽差。此外由于視蓋是魚類的視覺中樞[3,6,9,12—14], 有將刺激從視網(wǎng)膜傳遞入腦形成視覺, 同時(shí)還具有控制魚類位置和移動(dòng)的功能, 墨龍的圍腦室層厚度降低, 推測(cè)其游動(dòng)及平衡能力也較差。

此外, 與前人觀察結(jié)果相似[2], 也發(fā)現(xiàn)墨龍視網(wǎng)膜色素上皮層向桿錐層運(yùn)動(dòng)并交錯(cuò)對(duì)插, 推測(cè)是墨龍的色素上皮層可以防止因視網(wǎng)膜面積增大而導(dǎo)致桿錐層產(chǎn)生強(qiáng)烈光感。

總之, 人工選擇不但改變了金魚的外形, 而且使得金魚的中樞神經(jīng)組織結(jié)構(gòu)也發(fā)生了較大變化,墨龍的眼徑較紅鯽大, 視網(wǎng)膜面積也較大, 導(dǎo)致傳遞神經(jīng)沖動(dòng)的神經(jīng)纖維和神經(jīng)元增多。推測(cè)墨龍視網(wǎng)膜色素上皮層向桿錐層運(yùn)動(dòng)并交錯(cuò), 外核層、神經(jīng)節(jié)細(xì)胞層厚度降低, 并且視蓋中處理信息的圍腦室層厚度也降低, 可以減少因視網(wǎng)膜面積增大而產(chǎn)生的強(qiáng)烈光感。

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A NEUROANATOMICAL STUDY OF THE BLACK DRAGONEYE’S RETINA AND OPTIC TECTUM IN COMPARISON WITH THE RED CRUCIAN’S

WAN An1,2, AN Shu-Qing1and HUA Tian-Miao3
(1. School of Life Science, Nanjing University, Nanjing 210093, China; 2. School of Resources and environment, Anqing Normal University, Anqing 246000, China; 3. School of Life Science, Anhui Normal University, Wuhu 241000, China)

The black dragoneye, a kind of dragoneye goldfish (Carassius auratus), has evolved from the red crucian. Four black dragoneyes and four red crucians, each being 10—12 cm long and weighing 35 g, were taken randomly and dissected. Their brains and eye-balls were observed using routine paraffin section and HE staining. The microscopic structures of the retinas and optic tectums of black dragoneye and the red crucian were studied under an optical microscope. We found that, compared with the red crucian, the retina of black dragoneye was 29.9% thinner, of which the outer plexiform layer increased 2.5%, and the inner plexiform layer increased 11.8%, while the inner nuclear layer decreased 21.6%, and the outer nuclear layer decreased 35.6%. Besides, the ganglion cell layer and the photoreceptor layer were thinner and irregular. However, optic tectum in black dragoneye was 28.9% thicker than that of the red crucian, except that the stratum periventriculare layer decreased 22.6%, all layers were incrassate, with the stratum album central layer increased 12.8%, the stratum griseum central layer increased 30.6%, the stratum fibrosum et griseum superficiale layer whose fibers were 21.9% denser than that of the red crusian, the stratum opticum layer was 91.7% thicker and the stratum marginal layer was 35.6% thicker. The results showed that the central nervous system of black dragoneye varied widely due to artificial selection and breeding. It can be inferred that the larger eyeball diameter and retina area of the black dragoneye resulted in an increase of fibers transmitted from the retina to the optic tectum, which probably was the main cause for the thicker layers of neurons and nerve fibers that transmit nervous impulse. To reduce the damage caused by strong light, the pigment epithelium layer moved to the photoreceptor layer interlacedly, and the inner nuclear layer of and outer nuclear layer the retina as well as the stratum periventriculare layer in its optic tectum became thinner. In addition, the thickness of the stratum periventriculare layer decreased, which can help to explain why the black dragoneye is not so good at swimming and balancing as the red crucian.

Carassius auratus; The black dragoneye; The red crucian; Retina; Optic tectum

Q111.2+2

A

1000-3207(2013)01-0100-06

10.7541/2013.100

圖版Ⅰ Plate Ⅰ
1．紅鯽視網(wǎng)膜; 2. 墨龍視網(wǎng)膜; 3. 紅鯽視蓋; 4. 墨龍視蓋; 如圖1, 2所示, 紅鯽視網(wǎng)膜結(jié)構(gòu)清晰, 而墨龍視網(wǎng)膜的色素上皮層(PEL)向桿錐層(PL)運(yùn)動(dòng), 并交錯(cuò)對(duì)插, 與紅鯽相比, 墨龍視網(wǎng)膜外核層(ONL)、內(nèi)核層(INL)厚度降低, 外網(wǎng)狀層(OPL)和內(nèi)網(wǎng)狀層(IPL)厚度增加;如圖3, 4所示, 紅鯽和墨龍的視蓋各層結(jié)構(gòu)均清晰可辨, 與紅鯽相比, 墨龍圍腦室層(SPV)厚度降低, 其余各層厚度均有所增加
1. The Black Dragoneye’s Retina; 2. The Black Dragoneye’s Retina; 3. The Red Crucian’s Optic Tectum; 4. The Black Dragoneye’s Optic Tectum. As Picture 1 and Picture 2 show, the retina of red crucian presents a legible structure, but in the retina of black dragoneye the pigment epithelium layer (PEL) moves to the photoreceptor layer (PL) interlacedly. Besides, the inner nuclear layer (INL) and outer nuclear layer (ONL) of black dragoneye were thinner, while its outer plexiform layer (OPL) and inner plexiform layer (IPL) were thicker; As Picture 3 and Picture 4 show, the optic tectum of red crucian and black dragoneye are legible. Compared with the red crucian, the stratum periventriculare (SPV) layer in optic tectum was thinner in the black dragoneye, and other layers were thicker. PEL: 色素上皮層Pigment epithelium layer; PL: 桿錐層Photoreceptor layer; ONL: 外核層Outer nuclear layer; OPL:外網(wǎng)狀層Outer plexiform layer; INL: 內(nèi)核層Inner nuclear layer; IPL: 內(nèi)網(wǎng)狀層Inner plexiform layer; GCL: 神經(jīng)節(jié)細(xì)胞層Ganglion cell layer; SPV: 圍腦室層Stratum periventriculare; SAC: 中央纖維層Stratum album central; SGC: 中央細(xì)胞層Stratum griseum central; SFGS: 表面纖維層Stratum fibrosum et griseum superficiale; SO: 視神經(jīng)層Stratum opticum; SM: 邊緣層Stratum marginal

2011-12-27;

2012-10-17

國家自然科學(xué)基金(31171082); 南京大學(xué)重點(diǎn)學(xué)科經(jīng)費(fèi); 安慶師范學(xué)院碩士科研啟動(dòng)基金(No.044-K05000000195); 校級(jí)教研項(xiàng)目(No.044-J10004000024)資助

萬安(1980—), 男, 安徽安慶人; 博士研究生; 研究方向?yàn)樯鷳B(tài)學(xué)。E-mail: wanan@aqtc.edu.cn

安樹青(1963—), E-mail: anshq@nju.edu.cn