LBT095-097惠珍 Done on Jan 7’09
The order of corrected neuronal events may yet be different from the three orders discussed so far, although we are entering a realm in which we are almost entirely reduced t speculation. The reason for suspecting a neuronal firing order that is different from the order of motor events is the anatomy of the peripheral nerves that innervate the respective muscle. Figure 3.8 shows diagrammatically the course of some of the nerves that are relevant to this discussion. There is considerable differences in the length of these nerves; notice especially the circuitous course of the recurrent nerve which is more than three times conduction time of impulses does not merely depend upon the distance of peripheral nerve that must be traversed but also upon the diameter of the nerve fibers (the smaller the fiber. the slower the condition). It is possible to measure the average length of the peripheral nerves and also to make microscopic studies of cross sections of these nerves and to measure the size of the diameters of fibers. This has been done (Auriti,1954; Krmpotic, 1958 and 1959), and it has been found that nerves involved in speech muscle innervation vary considerably in their composition of a nerve in terms of a caliber spectrum (a statistical frequency distribution of diameter sizes). Table 3.4 shows a few figures taken from the work of Krmpotic, indicating that there is always some overlap in the distributions but that there are nevertheless, marked differences among the various nerves. It is interesting that the the longest nerve, the recurrent, has statically the smallest fibers which thus aggravates the timing problem, introducing delays from two independent factors.
相連的神經位元的排列可能還不會害目前所提到的三種排列方式不同,雖然我們一直想要減少推測的範圍。對於作用的神經細胞保持環一的態度,是因為他和其他運動的單位不同,這運動的單位是其他週邊單位的解剖學,會造成個別肌肉受神經的控制。3.8圖了顯示跟這研究討論有關的一些神經的方式的圖表。在這些細胞的長度有很大的不同;特別是有再生周期的細胞的習慣的繞行方式,會比其他受神經控制的下顎肌肉的細胞還要3倍。這樣的脈動運作方式並不只靠週邊必須掛越的神經的距離,還要看細胞的直徑(越小的細胞,越少的傳導運作方式)。要量週邊的細胞的平均長度是可能的,也可以用顯微鏡來做研究,來衡量細胞的直徑。(Auriti, 1954; Krmpotic, 1958 and 1959)則有做過這些,也發現了:在細胞的組成上,發音器官的肌肉受神經控制的狀況有很大的不同。對於神經的纖維組成的表現方式,用光普的方式來呈現,是很正常的( 一種統計學的方式)。3.4表則是從Krmpotic的數據而來,顯示出總是在分佈的地方有重複的地方,此外,在不同的細胞是有不同的。越長的細胞,他的反應週期,統計出來是越小的細胞作用的時間總計的問題,是受其他兩個獨立變數影響的狀況,是很有趣的。
Kirmpotic, who has been specially interested in the problem of differential innervation time, has computed so-called neuromuscular indexes for all major muscles involved in speech. These indexes are simply the ratio of the average length of the nerve to the mean size of fiber diameter. Since there is still some uncertainty about the physiological interpretation of these determinations, we need not be concerned here with the details. Suffice it ti say that the anatomy of the nerves suggests that innervation time for intrinsic laryngeal muscles may easily be up to 30 msec longer than innervation time for muscles in and around the oral cavity.
Kirmpotic ,對這種不同的反應時間的問題,感到特別有有趣,對於發音所用到到的肌肉,他寫了個神經肌肉的指示表。這個只是表只代表了平均的細胞直徑的神經長度的比例。因為還有很多對這重分類的生理學上的詮釋的狀況,還不是很確定,我們不需要對這細節太關心。如果說神經的解剖學能滿足的話,則表示類部聲帶的肌肉的受神經控制的時間,很經意的就達到30 msec, 比起裡面的肌肉和口腔器官的受神經支配的時間還長。
Considering now that some articulatory events may last as short a period as 20 msec, it becomes a reasonable assumption that the firing order in the brain stem may at times be different from the order of events occurring at the periphery.
想想看有些的發音方式直續得時間是短到20msec,這樣也比較能解釋:偶爾,在腦幹運作的時間,是和其他相關的反應時間不一的假設。
A concrete example will clarify the point. Let us take the words obtain and optimal. If we make spectrograms of these words and measure and durations of the first three phones, we get values such as these:
Duration of Duration of
First vowel the labial stop the aspiration
Obtain 90 msec 170 msec 50 msec
Optimal 110 msec 160 msec 20 msec
有個比較具體的例子能解釋這。就拿obtain , optimal來說,如果我們用光譜來測量以及測量前三個音的發音時間,我們會得到以下的結果:
Among the acoustic cues for the discussion of certain voiced and unvoiced stops are the duration of the preceding vowel and the duration of the silence during which the lips are closed. Our perceptual acuity is highly sensitized to these temoral factores and , by the same token, our motor coordination is precise enough to times articulation with sufficient presision to bring about these distinctions. In the previous example the duration of the first vowel differs by only 20 msec and that of the labial stops by only 10 msec, and in the second instant, the duration of the aspiration (the only part of the /t/ we hear)lasts only 20 msec. Recent investigation carried out at the Haskins Laboratories have shown that a great variety of phonetic distinctions are entirely dependent upon timing factors of onset, duration, and cessation of voice where magnitudes well below 20 msec are od the essence. (See also the time relationships in the data reported on by Liberman, Delattre, Cooper, 1952; Schatz, 1954; and Liberman, Delatrre, Gerstman, and Cooper, 1956)
在區別某些有聲和無聲的塞音的一些語音上的技巧中,就是在嘴唇閉起來的,含前面的母音發音的長短,和有聲音的長短。我們敏銳的認知,是對這些有感覺的,也對提他相似的方式有感覺的。同樣的,我們的協和運動方式也對發音方式,有精確感覺,能夠區別不同之處。在前面的第一個母音的持續時間的例子中,不同的是只有20msec,唇塞音側只有10 msec,在第二的例子,是送氣的差別。(我們只聽到/t/)持續了20 msec 。最近的Haskins Laboratories的研究則指出:語音上的不同,是因為起始音、發音時間、聲音的中斷等因素影響,造成了強度低於20 msec。(參造Liberman, Delattre, Cooper, 1952; Schatz, 1954; and Liberman, Delatrre, Gerstman, and Cooper, 1956 的作品)
It is clear that there must be considerable precision in timing if laryngeal mechanisms are to integrated with oral ones. IN addition, there are hundreds of muscular adjustments to be made every second (that is a new neuromuscular event every few milliseconds) from which we begin to see the magnitude of the timing-ordering problem.
如果說聲帶機制,是和口腔整合在一起, 一定有對反應時間大量的衡量的標準,此外,有很多肌肉,對每秒的反應的時間的問題所造成的更動的變數,(就是在每千分之一秒鐘,神經反應的時間)。
In view of the above we are hardly surprised that we may encounter patients with lesions in the central nervous system who have difficulty in keeping elements of speech and language in the right order. An important theoretical discussion on this subject was contributed by Jakobson and Halle(1956). Patients who have this problem speak very slowly and indicate to the examiner that speaking constitutes a great effort of concentration: the behave as if they had to “think of the right order.” Even so, they will constantly mix up individual sounds; is may become si, task may become taks, syllable may become syballel and they are very hard to understand.*Generally, they tend to anticipate sounds that should come at a later time. The difficulty may occur in every other word, since their rate of production is markedly slow down. The improper anticipation of sounds is usually nit the patient’s only problem of ordering. He will also show a pathological propensity for spoonerism, entire words and phrase s being switched around or produced in advance. The patient’s intention is frequently made known to the examiner because the patient is fully aware if his difficulty and can often make a fresh start and repeat the intended sentence once more and correctly . Ina sever cases the corrected sentence will show new mistakes.
從上面看來,我們很難不對會遇到的中有神經有機能損壞的病人很難不感到驚訝,這些病人都有說話的問題, 無法分清楚正確順序,也無法分別正確單位.。對這個問題, Jakobson and Halle(1956)的理論架構,是有幫助的。有語言障礙的病人,說話都很慢,也會對檢查員指出,講話時很難集中注意力:他們的表現就像是需思考「正確的順序」。此外,他們也常對語音的順序搞混;例如is 變成si, 變成taks, syllable 變成 syballel ,而且,他們都很難瞭解。* 病人都會絕得聲音應該晚一點出來。這狀況,在他們製造語言很慢時,是每一秒都會遇到,代表每一個字都有這樣的問題。精楚的對每個聲音都認知,並不是病人唯一的語言順序的問題。病人都呈有spoonerism(字音的無意互換現象)的病理傾向,通常都會把整個片語位置亂換,或是,提前說。對檢查員來說,他們都會預期這樣的狀況,因為病人本身都知道自己有語言的障礙,常常會重新開始,或是把句子在講一次,或是,再修改一次。在嚴重一點的病人,就算是重新修改的句子,也是錯的。
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[生物] translation LBT004-007 惠珍Translation (Done)
LBT004-007惠珍 updated on Jan 5’ 09 (Done)
Chapter 1, section II)
II. FORM AND FUNCTION IN ONTOGENY
個體發生學的行是與功能
There is evidence (1) that the issues of the brain and the rest of the body constitute an organic, interdependent unit; and (2) that organisms are not programmed for their behavior by an ex-machina force, but instead they develop a program ontogenetically together with nervous and nonnervous tissues.
有證據顯示(1)「大腦的細胞組織」和「身體的其他的部分」是組成生物體這互相依賴的有機體。(2)這有機體並不是以他行為,或是「大機器」的力量(ex-machina force)來編碼的(programmed)> 而是,生物體根據神經和分神經組織而發展出「個體發生的編碼原則」。
(1) Mutual influence in the development of Nervous and Other Tissue
(1) 在神經和其他細胞相互影響的發展
Let us first consider the developmental relationship between nervous and other tissue. Our discussion can be divided into (a) metabolic or trophic relationships and (b) nonmetabolic, particularly mechanical relationhships.
讓我們把神經和其他細胞發展中玵互相影響的關係,考慮進去。我們的討論可分為2部。(a) 新陳代謝的或是營養的 (b)非新陳代謝的或是特定的物質上的關係。
(a) Trophic Relationships. Nervous tissue stands in an intimate relationship to other tissue anatomically contiguous to it. This is shown most clearly by the essential role played by nerves in the process of regeneration. There are a number of studies available which indicate that regeneration of an entire limb in lower vertebrates (fish, lizards, urodele, salamander, larval anuran, and postmetamorphic frog) and probably also in invertebrates is dependent on the presence of nerves in the amputated stump (Singer, 1959, Gutmann, 1964).
(a) 營養的關係。神經細胞和其他的很自然且連續在一起的細胞組織有很密切的關係。而這很明顯的顯示出在「再生」這過程,神經扮演了很基本的角色。有大量的文獻指出:「在低等脊椎動物上再生出一個肋骨」,無脊椎的如:(魚、蜥蜴、火蜥蜴、幼體無尾目動物、後變態的青蛙,及可能一些依賴神經存在的斷尾的部分。)
In a series of experiments by Schotté and Butler (1944), Singer (1947), and their students, and Nicholas (1949), it has been shown that an amputated limb will not regenerate unless an intact nerve is either present in the remaining stump from the beginning or is transplanted into the cut surface by autograft (however, see Thornton and Steen, 1962). Morphogenesis, that is, an orderly sequence of tissue differentiation and development of the lost appendage, will not ordinarily take place in the absence of living nervous issue during the very first stage of regeneration. If all nerves are removed form this stump during the earliest period, mitotic activity is dramatically slowed down; eventually some small amount of connective tissue, cartilage, and muscle may form in a disorderly nonfunctional fashion, giving a shriveled and shapeless appearance to the stump. If viable nervous tissue is not present from the start of the amputation but brought into the so-called blastema shortly afterward, regeneration takes place but the regenerate limb is poorly developed. The nerve need not be present throughout the entire period of regeneration; once the limb has begun to grow and tissues are sufficiently differentiated, the nerve may be removed without impariment of the morphegenetic potency acquired by these tissues during their earliest stage of formation. Singer (1974) has shown that it does not matter for regeneration what type of nerve, whether motor, sensory, or autonomic, is present in the blastema. It is merely the amount of nervous tissue present that controls the regenerative possibilities. Apparently a product of nerve-cell metabolism induces morphogenesis in the blastema.
在Scotte和 Bluter , Singer 和他們的學生及Nicholas,一系列的實驗中,顯示出:被截斷的肋骨是不會再生,除非他的神經一開始就仍然在存在的斷腿; 或是移植到自體移植皮膚表面的東西。(但,看看Thronton & Steen) 器官再生,就是細胞不同化和很多附屬器官的表層一連串的過程,在很早期的再生狀況下,缺乏活體神經組織之下,是不會發生的。如果所有的義肢、斷腿,都在早期都被移除,那麼有可絲分裂,就會大量下降活動; 甚至其他一些相關連的組織、軟骨、肌肉,都可能形成沒有規則、沒有功能的形狀、沒有用的、沒有形狀的斷腿的外觀。如果,神經細胞不是在開始截肢的時候就找到,是被帶到之後所謂的胚芽,「再生」就會開始。但是再生的肋骨市是「不完全發育」,神經不會在整個再生過程中呈現出來; 一旦,肋骨開始生長,而且組織得到足夠的養分,神經就可以從器官器官再生的部位移除,力量,是細胞在早期形成的時間得到的Singer,則證明了:任何細胞組織,對再生來說,是不相關的,不管是運動、感覺、自動的,都在胚芽中呈現。只是在細胞組織中呈現,然後控制再生的可能性,顯然地,細胞組織新陳代謝的解果會包括批牙器官再生。
These studies leave many questions about the biochemistry of embryology and growth unanswered; yet they do give us a glimpse of the complete interdependence and the natural integration of different tissues in the animal body. This impression is further strengthened if we consider some of the other trophic relationships that nerves have to peripheral tissue (for instance, the well-known fact of denervation atrophy). If the axon of a motor neuron is cut, the portion distal of the cut will die promptly, presumably because of its separation from its source of supply of vital substances (Gerard, 1950). But this is not the extent of the degenerative changes following the section of a motor nerve. The muscle innervated by the nerve will also undergo dystrophic changes with an extremely characteristic histological appearance. The loss of muscle substance is not due to a “functional” disturbance, such as the inhibition of nerve impulse transmission (Hamburger & Levi-Montalcini, 1950) nor due to disuse of the muscle; the muscle cannot be saved from atrophy by passive exercise. Indeed, the metabolic interdependence of nonnervous, peripheral tissue and nervous tissue is proven by the fact that the nerves themselves must have anatomic continuity with muscles for proper metabolic function. Severance of nerve from muscle will induce retrograde changes in the body of the neuron (the soma), known as chromatolysis, which is a sign of dysfunction.
對於生物化學胚胎學成長而言,對很多問題, 很多研究都還沒有有蒙卻的解答。但是, 這與給我們對於完全的依賴 或是再不同動物種類、細胞之間的整合演變,
有一種簡單的了解. 這也亙加強了我們對於其他的發育的關係的關感:細胞只是附屬的(例如:有名的去掉植)【動】脈序;脈狀;翅脈的例子). 如果運動神經的軸索被切斷, 末端的組織就會因為和濫以維生的物質分開而開始死亡。(Gerard1950) 但不是因為退化的物質, 改變運動神經. 而是受受神經支配的 肌肉會被大量的組織物質污染. 這種肌肉的損失是因為「功能亂流」例如;脈搏的跳動(Hamburger1950) 也不是因為肌肉組織造成的 ; 肌肉無法被動的發育停止. 反而是, 非細胞間的心陳代謝作用, 其他週邊的組織和神經細胞被證明了:是自動連續性的神經而且帶來了合適的新陳代謝. 有很多的肌肉神經, 會在神經元內減少, (體細胞) 就是「色原溶解;核染質溶解;染色質溶解消失;色素溶解消失」也是代表了「官能障礙」
Perhaps the most striking evidence for the subtle but definite interdependence of peripheral structures and the central nervous system is provided by the stunted growth resulting from large cerebro-hemispheric and specifically parietal lobe lesions in the neonate human. This phenomenon was described by Macdonald Critchley (1955) and has been generally known to occur in connection with a condition called infantile hemiplegia. The stunting of the body side contralateral to the brain lesion occurs both in congenital and in acquired infantile hemiplegia and is not due to disuse, first because all tissues in the extremities involved are equally affected and second, because the arrest of growth starts at birth in the congenital cases, that is, before either the affected or the unaffected side is actively being used (Holt, 1961). These cases are even more interesting in the present context than the denervation atrohy, because here we see a relation between the highest level of the central nervous system and nonnervous structures of the periphery. Because the cells of the cerebral cortex are separated from those on the periphery by several internuncial neurons, there is an indication of a very subtle control that the higher centers of the central nervous system appear to exe3rcise upon the development of the body as a whole.
也許最驚人的事是:精細但與其他的結構相連,和由神奇的成長的中央神經系統
,在未滿月的)嬰兒時,會造成腦部的和顱頂骨腦的葉的器官損害。Macdonald 曾提過這種現象,而且他也想知道這種關聯:「未滿月的」嬰兒的未滿月的)嬰兒」。這種一連串的對於大腦的損害也會發生在先天的和後來得到的嬰兒的半身不遂,也不全是因為廢棄不使用的關係,而是因為末端的組織已經被感染。再者,在先天的生長案例中,還沒被感染的也會污染,這種狀況常常發生(Holt 1961)這狀況比起沒有脈落的萎縮現在的內容還有趣。因為我們看到了更高一樓的中央神經系統和其他的附近非神經的結構。因為大腦的皮質細胞是由不同的(神經細胞間之)聯絡的相連而成, 所以,更高一樓的中央神經系統是很精密的活動著,這也反映在整體的身體運作。
(b) Mechanical Relationships. In addition to the metabolic influences between nervous tissue and other tissue, morphogenesis is controlled by several other factors, some of which are indirectly related to neurophysiology. A good illustration is furnished by the mechanical forces exerted upon growing tissue, particularly bone, which stimulate cell division in certain directions. As muscles are innervated and begin to function, they exert a pull upon the boned to which they are attached and thus help shape the internal structure of this tissue.
(b) 機械學的,力學的;物理的關係
除了介於在神經組織和其他組織中的新陳代謝作用的影響之外,器官再生控制了幾個重要的因素,有些和神經生理學並沒有直接的關聯。作用暫成長的組織上的物理上力量,就是個好例子,特別是骨頭,在某種程度會刺激細胞分化。當肌肉該使受神經約束,該使運作時,會運作在附著的骨頭上,幫助使其形成內部組織。
It has often been noted (and, unfortunately, frequently been emphasized out of all proportion to its true importance) that the architecture of certain organs is ideally suited to their function. An excellent example of this was provided by D’Arcy Thompson (1942) who wrote, “In all the mechanical side of anatomy nothing can be more beautiful than the construction of a vulture’s metacarpal bone. The engineer sees in it a perfect Warren’s truss, just such a one as is often used for a main rib in an aeroplane.” The fundamental schema of the shape of individual bones and the skeleton as a whole are undoubtedly the result of evolutionary processes including selection and adaptation on a phylogenetic rather than an ontogenetic scale (Hackenbroch, 1957-1962). However, the actual realization of what is only potentially present in the fertilized egg is largely dependent upon factors which are active during ontogenesis. This is vest illustrated by the development of the internal structure of bones.
(很不幸的,常很少對於其重要性有成相對應的重視,有些器官的組成就是很符合本身的功能 )是很常被注意的。D’Arcy Thompson 就提供了個很好的例子,他寫道:『在解剖學的物理現象中,謀有什麼比禿鷹的手掌的骨頭更漂亮。這位工程師把它當成「養兔場的兔子」疝帶,托帶(Warren’s truss),就像飛機的主要肋骨架構』每跟骨頭的形狀的基本形狀都不同,而且,整個身體的支架也是進化的結果,包含了動植物種類史的選擇和突變,而不是個體發生的方向(Hachkenbroch,1957-1962)。但是,真正有可能在受精卵出現的時候,適一存在個體發生的過程中。骨頭的內部結構正好說明了這一點。
In 1866, a Swiss engineer, Culmann, noted that the internal trusses in the head of a human femur, anatomically known as trabeculae, were oriented in exactly the direction of the lines of maximum internal stress. He drew a diagram of a curved rod showing the lines of stress resulting from the application of a load from above (Fig. 1.1a). The model somewhat resembled the head of a derrick which he had just designed, and it is therefore referred to as a crane’s head. It bears a striking similarity to a section of the head of the femur (Fig. 1.1c). Culmann’s idea gave rise to J. Wolff’s famous theory expressed in his monograph on The Law of Bone Transformation (1870), according to which every change in the function of a bone produces changes in its trabecular architecture and external form in conformity with mathematical, static laws. The theory has since been criticized (Küntsher, 1934, 1936), modified (Murray, 1936), and elaborated upon (Evans, 1955, 1957, 1960; Carey, 1929). However, the idea is widely accepted today that muscles, through the tonus already present on early embryonic life, exert essential forces upon the growing bone. These pressures, together with those produced by differential growth of various parts of the embryo, result in stresses and strains which are the prerequisite stimuli for proper bone formation. [end]
在1866年,一個瑞士的工程師,Columann則對於人體的股骨的街頭內部分隔帶,則會在最大的內部壓力下,精確的呈現了其線條。他劃一個灣區的桿,則顯示壓力線下壓力會造成上面所提到的作用(Fig. 1.1a)。這種模式就好像他設計的油井的鐵支架一樣,也像個鶴的頭。它可以接受股骨的接頭部份所承受的壓力(Fig. 1.1c)。Culmann的想法影響了後來的J. Wolff有名的理論,顯現在他1870《骨頭的轉變法則》專題論文中 :在製造每個骨頭的功能時都會改變他的分隔帶的狀況,而且,外部形式時複合數學、統計法則的。(Kuntscher 1934,1936), 攻擊得理論,(Murry, 1936) 修改他,(Evans, 1955, 1957,1960; Carey, 1929)則把這理論發揚光大。但是,在今天大部人接受的肌肉理論,是藉由早期的初期/胎兒生物所呈現出肌肉強直性,在成長中的骨頭運作其基本的力量。這些壓力伴隨著胚胎不同階段的成長過程,會造成壓力,及對骨頭成長不可或缺的刺激的拉緊。
l New Words: Ontogeny: 【生】個體發生;個體發生學 Independent : 相互依賴的;互助的 ontogenetically: adj of ontogeny Ex-machina : is an American creator-owned comic book series created by Brian K. Vaughan and Tony Harris, and published by DC Comics under the Wildstorm imprint. Metabolic: 新陳代謝的 Trophic營養的;有關營養的 Regeneration:再生 contiguous連續的 limb 肢;臂;腳;翼; 大樹枝,主枝; 分支;執行者,代理人 vertebrates 脊椎動物 lizard: 蜥蜴 urodele salamander (傳說中生活在火中的)火蜥蜴,火蛇;火精,火怪 larval anuran, 幼蟲的;幼體的 anuran無尾目動物(蛙、蛤蟆等等) postmetamorphic :後變態的 amputated 醫】切斷;鋸掉;截(肢) stump口】腿;義肢[P] autograft 醫】自身移植物(如皮膚等) Morphogenesis, 器官發生 appendage, 生】附屬器官,附屬肢體(如臂、腿、尾等) mitotic 生】有絲分裂的 cartilage 【解】軟骨原骨,軟骨成骨 shriveled使束手無策;使無能為力;使無用 blastema 【生】胚芽 autonomic【生】自律的 embryology 胚胎學 denervation;去掉植】【動】脈序;脈狀;翅脈atrophy醫】萎縮;發育停止;虛脫 distal 【解】末梢部的;末端的 innervated使(器官、肌肉等)受神經支配 dystrophic (家庭或工業垃圾等對水的)污染 histological 組織學的 retrograde後退的;逆行的;退化的 soma【生】體細胞 chromatolysis【生】【醫】色原溶解;核染質溶解;染色質溶解消失;色素溶解消失 dysfunction. 官能不良;官能障礙
l cerebro-hemispheric 腦部的parietal【生】腔壁的;【解】顱頂骨的;【植】子房壁的 lobe【解】(腦、 肺等的)葉 lesions機能障礙;器官損害neonate (未滿月的)嬰兒 hemiplegia半身麻痺;半身不遂contralateral congenital天生的;先天的
l nternuncial 【解】(神經細胞間之)聯絡的 furnished 給(房間)配置(傢俱等);裝備[(+with)] metacarpal【解】掌的;掌部的 Warren’s truss Aeroplane【英】飛機[C] Femur解】股骨;【昆】股節 Trabeculae【解】分隔帶
l derrick:【機】轉臂起重機 / 油井的鐵架塔embryonic 胚芽的;胎兒的 prerequisite不可缺的;事先需要的;必修
LBT004-007惠珍 updated on Jan 5’ 09 (Done)
Chapter 1, section II)
II. FORM AND FUNCTION IN ONTOGENY
個體發生學的行是與功能
There is evidence (1) that the issues of the brain and the rest of the body constitute an organic, interdependent unit; and (2) that organisms are not programmed for their behavior by an ex-machina force, but instead they develop a program ontogenetically together with nervous and nonnervous tissues.
有證據顯示(1)「大腦的細胞組織」和「身體的其他的部分」是組成生物體這互相依賴的有機體。(2)這有機體並不是以他行為,或是「大機器」的力量(ex-machina force)來編碼的(programmed)> 而是,生物體根據神經和分神經組織而發展出「個體發生的編碼原則」。
(1) Mutual influence in the development of Nervous and Other Tissue
(1) 在神經和其他細胞相互影響的發展
Let us first consider the developmental relationship between nervous and other tissue. Our discussion can be divided into (a) metabolic or trophic relationships and (b) nonmetabolic, particularly mechanical relationhships.
讓我們把神經和其他細胞發展中玵互相影響的關係,考慮進去。我們的討論可分為2部。(a) 新陳代謝的或是營養的 (b)非新陳代謝的或是特定的物質上的關係。
(a) Trophic Relationships. Nervous tissue stands in an intimate relationship to other tissue anatomically contiguous to it. This is shown most clearly by the essential role played by nerves in the process of regeneration. There are a number of studies available which indicate that regeneration of an entire limb in lower vertebrates (fish, lizards, urodele, salamander, larval anuran, and postmetamorphic frog) and probably also in invertebrates is dependent on the presence of nerves in the amputated stump (Singer, 1959, Gutmann, 1964).
(a) 營養的關係。神經細胞和其他的很自然且連續在一起的細胞組織有很密切的關係。而這很明顯的顯示出在「再生」這過程,神經扮演了很基本的角色。有大量的文獻指出:「在低等脊椎動物上再生出一個肋骨」,無脊椎的如:(魚、蜥蜴、火蜥蜴、幼體無尾目動物、後變態的青蛙,及可能一些依賴神經存在的斷尾的部分。)
In a series of experiments by Schotté and Butler (1944), Singer (1947), and their students, and Nicholas (1949), it has been shown that an amputated limb will not regenerate unless an intact nerve is either present in the remaining stump from the beginning or is transplanted into the cut surface by autograft (however, see Thornton and Steen, 1962). Morphogenesis, that is, an orderly sequence of tissue differentiation and development of the lost appendage, will not ordinarily take place in the absence of living nervous issue during the very first stage of regeneration. If all nerves are removed form this stump during the earliest period, mitotic activity is dramatically slowed down; eventually some small amount of connective tissue, cartilage, and muscle may form in a disorderly nonfunctional fashion, giving a shriveled and shapeless appearance to the stump. If viable nervous tissue is not present from the start of the amputation but brought into the so-called blastema shortly afterward, regeneration takes place but the regenerate limb is poorly developed. The nerve need not be present throughout the entire period of regeneration; once the limb has begun to grow and tissues are sufficiently differentiated, the nerve may be removed without impariment of the morphegenetic potency acquired by these tissues during their earliest stage of formation. Singer (1974) has shown that it does not matter for regeneration what type of nerve, whether motor, sensory, or autonomic, is present in the blastema. It is merely the amount of nervous tissue present that controls the regenerative possibilities. Apparently a product of nerve-cell metabolism induces morphogenesis in the blastema.
在Scotte和 Bluter , Singer 和他們的學生及Nicholas,一系列的實驗中,顯示出:被截斷的肋骨是不會再生,除非他的神經一開始就仍然在存在的斷腿; 或是移植到自體移植皮膚表面的東西。(但,看看Thronton & Steen) 器官再生,就是細胞不同化和很多附屬器官的表層一連串的過程,在很早期的再生狀況下,缺乏活體神經組織之下,是不會發生的。如果所有的義肢、斷腿,都在早期都被移除,那麼有可絲分裂,就會大量下降活動; 甚至其他一些相關連的組織、軟骨、肌肉,都可能形成沒有規則、沒有功能的形狀、沒有用的、沒有形狀的斷腿的外觀。如果,神經細胞不是在開始截肢的時候就找到,是被帶到之後所謂的胚芽,「再生」就會開始。但是再生的肋骨市是「不完全發育」,神經不會在整個再生過程中呈現出來; 一旦,肋骨開始生長,而且組織得到足夠的養分,神經就可以從器官器官再生的部位移除,力量,是細胞在早期形成的時間得到的Singer,則證明了:任何細胞組織,對再生來說,是不相關的,不管是運動、感覺、自動的,都在胚芽中呈現。只是在細胞組織中呈現,然後控制再生的可能性,顯然地,細胞組織新陳代謝的解果會包括批牙器官再生。
These studies leave many questions about the biochemistry of embryology and growth unanswered; yet they do give us a glimpse of the complete interdependence and the natural integration of different tissues in the animal body. This impression is further strengthened if we consider some of the other trophic relationships that nerves have to peripheral tissue (for instance, the well-known fact of denervation atrophy). If the axon of a motor neuron is cut, the portion distal of the cut will die promptly, presumably because of its separation from its source of supply of vital substances (Gerard, 1950). But this is not the extent of the degenerative changes following the section of a motor nerve. The muscle innervated by the nerve will also undergo dystrophic changes with an extremely characteristic histological appearance. The loss of muscle substance is not due to a “functional” disturbance, such as the inhibition of nerve impulse transmission (Hamburger & Levi-Montalcini, 1950) nor due to disuse of the muscle; the muscle cannot be saved from atrophy by passive exercise. Indeed, the metabolic interdependence of nonnervous, peripheral tissue and nervous tissue is proven by the fact that the nerves themselves must have anatomic continuity with muscles for proper metabolic function. Severance of nerve from muscle will induce retrograde changes in the body of the neuron (the soma), known as chromatolysis, which is a sign of dysfunction.
對於生物化學胚胎學成長而言,對很多問題, 很多研究都還沒有有蒙卻的解答。但是, 這與給我們對於完全的依賴 或是再不同動物種類、細胞之間的整合演變,
有一種簡單的了解. 這也亙加強了我們對於其他的發育的關係的關感:細胞只是附屬的(例如:有名的去掉植)【動】脈序;脈狀;翅脈的例子). 如果運動神經的軸索被切斷, 末端的組織就會因為和濫以維生的物質分開而開始死亡。(Gerard1950) 但不是因為退化的物質, 改變運動神經. 而是受受神經支配的 肌肉會被大量的組織物質污染. 這種肌肉的損失是因為「功能亂流」例如;脈搏的跳動(Hamburger1950) 也不是因為肌肉組織造成的 ; 肌肉無法被動的發育停止. 反而是, 非細胞間的心陳代謝作用, 其他週邊的組織和神經細胞被證明了:是自動連續性的神經而且帶來了合適的新陳代謝. 有很多的肌肉神經, 會在神經元內減少, (體細胞) 就是「色原溶解;核染質溶解;染色質溶解消失;色素溶解消失」也是代表了「官能障礙」
Perhaps the most striking evidence for the subtle but definite interdependence of peripheral structures and the central nervous system is provided by the stunted growth resulting from large cerebro-hemispheric and specifically parietal lobe lesions in the neonate human. This phenomenon was described by Macdonald Critchley (1955) and has been generally known to occur in connection with a condition called infantile hemiplegia. The stunting of the body side contralateral to the brain lesion occurs both in congenital and in acquired infantile hemiplegia and is not due to disuse, first because all tissues in the extremities involved are equally affected and second, because the arrest of growth starts at birth in the congenital cases, that is, before either the affected or the unaffected side is actively being used (Holt, 1961). These cases are even more interesting in the present context than the denervation atrohy, because here we see a relation between the highest level of the central nervous system and nonnervous structures of the periphery. Because the cells of the cerebral cortex are separated from those on the periphery by several internuncial neurons, there is an indication of a very subtle control that the higher centers of the central nervous system appear to exe3rcise upon the development of the body as a whole.
也許最驚人的事是:精細但與其他的結構相連,和由神奇的成長的中央神經系統
,在未滿月的)嬰兒時,會造成腦部的和顱頂骨腦的葉的器官損害。Macdonald 曾提過這種現象,而且他也想知道這種關聯:「未滿月的」嬰兒的未滿月的)嬰兒」。這種一連串的對於大腦的損害也會發生在先天的和後來得到的嬰兒的半身不遂,也不全是因為廢棄不使用的關係,而是因為末端的組織已經被感染。再者,在先天的生長案例中,還沒被感染的也會污染,這種狀況常常發生(Holt 1961)這狀況比起沒有脈落的萎縮現在的內容還有趣。因為我們看到了更高一樓的中央神經系統和其他的附近非神經的結構。因為大腦的皮質細胞是由不同的(神經細胞間之)聯絡的相連而成, 所以,更高一樓的中央神經系統是很精密的活動著,這也反映在整體的身體運作。
(b) Mechanical Relationships. In addition to the metabolic influences between nervous tissue and other tissue, morphogenesis is controlled by several other factors, some of which are indirectly related to neurophysiology. A good illustration is furnished by the mechanical forces exerted upon growing tissue, particularly bone, which stimulate cell division in certain directions. As muscles are innervated and begin to function, they exert a pull upon the boned to which they are attached and thus help shape the internal structure of this tissue.
(b) 機械學的,力學的;物理的關係
除了介於在神經組織和其他組織中的新陳代謝作用的影響之外,器官再生控制了幾個重要的因素,有些和神經生理學並沒有直接的關聯。作用暫成長的組織上的物理上力量,就是個好例子,特別是骨頭,在某種程度會刺激細胞分化。當肌肉該使受神經約束,該使運作時,會運作在附著的骨頭上,幫助使其形成內部組織。
It has often been noted (and, unfortunately, frequently been emphasized out of all proportion to its true importance) that the architecture of certain organs is ideally suited to their function. An excellent example of this was provided by D’Arcy Thompson (1942) who wrote, “In all the mechanical side of anatomy nothing can be more beautiful than the construction of a vulture’s metacarpal bone. The engineer sees in it a perfect Warren’s truss, just such a one as is often used for a main rib in an aeroplane.” The fundamental schema of the shape of individual bones and the skeleton as a whole are undoubtedly the result of evolutionary processes including selection and adaptation on a phylogenetic rather than an ontogenetic scale (Hackenbroch, 1957-1962). However, the actual realization of what is only potentially present in the fertilized egg is largely dependent upon factors which are active during ontogenesis. This is vest illustrated by the development of the internal structure of bones.
(很不幸的,常很少對於其重要性有成相對應的重視,有些器官的組成就是很符合本身的功能 )是很常被注意的。D’Arcy Thompson 就提供了個很好的例子,他寫道:『在解剖學的物理現象中,謀有什麼比禿鷹的手掌的骨頭更漂亮。這位工程師把它當成「養兔場的兔子」疝帶,托帶(Warren’s truss),就像飛機的主要肋骨架構』每跟骨頭的形狀的基本形狀都不同,而且,整個身體的支架也是進化的結果,包含了動植物種類史的選擇和突變,而不是個體發生的方向(Hachkenbroch,1957-1962)。但是,真正有可能在受精卵出現的時候,適一存在個體發生的過程中。骨頭的內部結構正好說明了這一點。
In 1866, a Swiss engineer, Culmann, noted that the internal trusses in the head of a human femur, anatomically known as trabeculae, were oriented in exactly the direction of the lines of maximum internal stress. He drew a diagram of a curved rod showing the lines of stress resulting from the application of a load from above (Fig. 1.1a). The model somewhat resembled the head of a derrick which he had just designed, and it is therefore referred to as a crane’s head. It bears a striking similarity to a section of the head of the femur (Fig. 1.1c). Culmann’s idea gave rise to J. Wolff’s famous theory expressed in his monograph on The Law of Bone Transformation (1870), according to which every change in the function of a bone produces changes in its trabecular architecture and external form in conformity with mathematical, static laws. The theory has since been criticized (Küntsher, 1934, 1936), modified (Murray, 1936), and elaborated upon (Evans, 1955, 1957, 1960; Carey, 1929). However, the idea is widely accepted today that muscles, through the tonus already present on early embryonic life, exert essential forces upon the growing bone. These pressures, together with those produced by differential growth of various parts of the embryo, result in stresses and strains which are the prerequisite stimuli for proper bone formation. [end]
在1866年,一個瑞士的工程師,Columann則對於人體的股骨的街頭內部分隔帶,則會在最大的內部壓力下,精確的呈現了其線條。他劃一個灣區的桿,則顯示壓力線下壓力會造成上面所提到的作用(Fig. 1.1a)。這種模式就好像他設計的油井的鐵支架一樣,也像個鶴的頭。它可以接受股骨的接頭部份所承受的壓力(Fig. 1.1c)。Culmann的想法影響了後來的J. Wolff有名的理論,顯現在他1870《骨頭的轉變法則》專題論文中 :在製造每個骨頭的功能時都會改變他的分隔帶的狀況,而且,外部形式時複合數學、統計法則的。(Kuntscher 1934,1936), 攻擊得理論,(Murry, 1936) 修改他,(Evans, 1955, 1957,1960; Carey, 1929)則把這理論發揚光大。但是,在今天大部人接受的肌肉理論,是藉由早期的初期/胎兒生物所呈現出肌肉強直性,在成長中的骨頭運作其基本的力量。這些壓力伴隨著胚胎不同階段的成長過程,會造成壓力,及對骨頭成長不可或缺的刺激的拉緊。
l New Words: Ontogeny: 【生】個體發生;個體發生學 Independent : 相互依賴的;互助的 ontogenetically: adj of ontogeny Ex-machina : is an American creator-owned comic book series created by Brian K. Vaughan and Tony Harris, and published by DC Comics under the Wildstorm imprint. Metabolic: 新陳代謝的 Trophic營養的;有關營養的 Regeneration:再生 contiguous連續的 limb 肢;臂;腳;翼; 大樹枝,主枝; 分支;執行者,代理人 vertebrates 脊椎動物 lizard: 蜥蜴 urodele salamander (傳說中生活在火中的)火蜥蜴,火蛇;火精,火怪 larval anuran, 幼蟲的;幼體的 anuran無尾目動物(蛙、蛤蟆等等) postmetamorphic :後變態的 amputated 醫】切斷;鋸掉;截(肢) stump口】腿;義肢[P] autograft 醫】自身移植物(如皮膚等) Morphogenesis, 器官發生 appendage, 生】附屬器官,附屬肢體(如臂、腿、尾等) mitotic 生】有絲分裂的 cartilage 【解】軟骨原骨,軟骨成骨 shriveled使束手無策;使無能為力;使無用 blastema 【生】胚芽 autonomic【生】自律的 embryology 胚胎學 denervation;去掉植】【動】脈序;脈狀;翅脈atrophy醫】萎縮;發育停止;虛脫 distal 【解】末梢部的;末端的 innervated使(器官、肌肉等)受神經支配 dystrophic (家庭或工業垃圾等對水的)污染 histological 組織學的 retrograde後退的;逆行的;退化的 soma【生】體細胞 chromatolysis【生】【醫】色原溶解;核染質溶解;染色質溶解消失;色素溶解消失 dysfunction. 官能不良;官能障礙
l cerebro-hemispheric 腦部的parietal【生】腔壁的;【解】顱頂骨的;【植】子房壁的 lobe【解】(腦、 肺等的)葉 lesions機能障礙;器官損害neonate (未滿月的)嬰兒 hemiplegia半身麻痺;半身不遂contralateral congenital天生的;先天的
l nternuncial 【解】(神經細胞間之)聯絡的 furnished 給(房間)配置(傢俱等);裝備[(+with)] metacarpal【解】掌的;掌部的 Warren’s truss Aeroplane【英】飛機[C] Femur解】股骨;【昆】股節 Trabeculae【解】分隔帶
l derrick:【機】轉臂起重機 / 油井的鐵架塔embryonic 胚芽的;胎兒的 prerequisite不可缺的;事先需要的;必修
Chapter 1, section II)
II. FORM AND FUNCTION IN ONTOGENY
個體發生學的行是與功能
There is evidence (1) that the issues of the brain and the rest of the body constitute an organic, interdependent unit; and (2) that organisms are not programmed for their behavior by an ex-machina force, but instead they develop a program ontogenetically together with nervous and nonnervous tissues.
有證據顯示(1)「大腦的細胞組織」和「身體的其他的部分」是組成生物體這互相依賴的有機體。(2)這有機體並不是以他行為,或是「大機器」的力量(ex-machina force)來編碼的(programmed)> 而是,生物體根據神經和分神經組織而發展出「個體發生的編碼原則」。
(1) Mutual influence in the development of Nervous and Other Tissue
(1) 在神經和其他細胞相互影響的發展
Let us first consider the developmental relationship between nervous and other tissue. Our discussion can be divided into (a) metabolic or trophic relationships and (b) nonmetabolic, particularly mechanical relationhships.
讓我們把神經和其他細胞發展中玵互相影響的關係,考慮進去。我們的討論可分為2部。(a) 新陳代謝的或是營養的 (b)非新陳代謝的或是特定的物質上的關係。
(a) Trophic Relationships. Nervous tissue stands in an intimate relationship to other tissue anatomically contiguous to it. This is shown most clearly by the essential role played by nerves in the process of regeneration. There are a number of studies available which indicate that regeneration of an entire limb in lower vertebrates (fish, lizards, urodele, salamander, larval anuran, and postmetamorphic frog) and probably also in invertebrates is dependent on the presence of nerves in the amputated stump (Singer, 1959, Gutmann, 1964).
(a) 營養的關係。神經細胞和其他的很自然且連續在一起的細胞組織有很密切的關係。而這很明顯的顯示出在「再生」這過程,神經扮演了很基本的角色。有大量的文獻指出:「在低等脊椎動物上再生出一個肋骨」,無脊椎的如:(魚、蜥蜴、火蜥蜴、幼體無尾目動物、後變態的青蛙,及可能一些依賴神經存在的斷尾的部分。)
In a series of experiments by Schotté and Butler (1944), Singer (1947), and their students, and Nicholas (1949), it has been shown that an amputated limb will not regenerate unless an intact nerve is either present in the remaining stump from the beginning or is transplanted into the cut surface by autograft (however, see Thornton and Steen, 1962). Morphogenesis, that is, an orderly sequence of tissue differentiation and development of the lost appendage, will not ordinarily take place in the absence of living nervous issue during the very first stage of regeneration. If all nerves are removed form this stump during the earliest period, mitotic activity is dramatically slowed down; eventually some small amount of connective tissue, cartilage, and muscle may form in a disorderly nonfunctional fashion, giving a shriveled and shapeless appearance to the stump. If viable nervous tissue is not present from the start of the amputation but brought into the so-called blastema shortly afterward, regeneration takes place but the regenerate limb is poorly developed. The nerve need not be present throughout the entire period of regeneration; once the limb has begun to grow and tissues are sufficiently differentiated, the nerve may be removed without impariment of the morphegenetic potency acquired by these tissues during their earliest stage of formation. Singer (1974) has shown that it does not matter for regeneration what type of nerve, whether motor, sensory, or autonomic, is present in the blastema. It is merely the amount of nervous tissue present that controls the regenerative possibilities. Apparently a product of nerve-cell metabolism induces morphogenesis in the blastema.
在Scotte和 Bluter , Singer 和他們的學生及Nicholas,一系列的實驗中,顯示出:被截斷的肋骨是不會再生,除非他的神經一開始就仍然在存在的斷腿; 或是移植到自體移植皮膚表面的東西。(但,看看Thronton & Steen) 器官再生,就是細胞不同化和很多附屬器官的表層一連串的過程,在很早期的再生狀況下,缺乏活體神經組織之下,是不會發生的。如果所有的義肢、斷腿,都在早期都被移除,那麼有可絲分裂,就會大量下降活動; 甚至其他一些相關連的組織、軟骨、肌肉,都可能形成沒有規則、沒有功能的形狀、沒有用的、沒有形狀的斷腿的外觀。如果,神經細胞不是在開始截肢的時候就找到,是被帶到之後所謂的胚芽,「再生」就會開始。但是再生的肋骨市是「不完全發育」,神經不會在整個再生過程中呈現出來; 一旦,肋骨開始生長,而且組織得到足夠的養分,神經就可以從器官器官再生的部位移除,力量,是細胞在早期形成的時間得到的Singer,則證明了:任何細胞組織,對再生來說,是不相關的,不管是運動、感覺、自動的,都在胚芽中呈現。只是在細胞組織中呈現,然後控制再生的可能性,顯然地,細胞組織新陳代謝的解果會包括批牙器官再生。
These studies leave many questions about the biochemistry of embryology and growth unanswered; yet they do give us a glimpse of the complete interdependence and the natural integration of different tissues in the animal body. This impression is further strengthened if we consider some of the other trophic relationships that nerves have to peripheral tissue (for instance, the well-known fact of denervation atrophy). If the axon of a motor neuron is cut, the portion distal of the cut will die promptly, presumably because of its separation from its source of supply of vital substances (Gerard, 1950). But this is not the extent of the degenerative changes following the section of a motor nerve. The muscle innervated by the nerve will also undergo dystrophic changes with an extremely characteristic histological appearance. The loss of muscle substance is not due to a “functional” disturbance, such as the inhibition of nerve impulse transmission (Hamburger & Levi-Montalcini, 1950) nor due to disuse of the muscle; the muscle cannot be saved from atrophy by passive exercise. Indeed, the metabolic interdependence of nonnervous, peripheral tissue and nervous tissue is proven by the fact that the nerves themselves must have anatomic continuity with muscles for proper metabolic function. Severance of nerve from muscle will induce retrograde changes in the body of the neuron (the soma), known as chromatolysis, which is a sign of dysfunction.
對於生物化學胚胎學成長而言,對很多問題, 很多研究都還沒有有蒙卻的解答。但是, 這與給我們對於完全的依賴 或是再不同動物種類、細胞之間的整合演變,
有一種簡單的了解. 這也亙加強了我們對於其他的發育的關係的關感:細胞只是附屬的(例如:有名的去掉植)【動】脈序;脈狀;翅脈的例子). 如果運動神經的軸索被切斷, 末端的組織就會因為和濫以維生的物質分開而開始死亡。(Gerard1950) 但不是因為退化的物質, 改變運動神經. 而是受受神經支配的 肌肉會被大量的組織物質污染. 這種肌肉的損失是因為「功能亂流」例如;脈搏的跳動(Hamburger1950) 也不是因為肌肉組織造成的 ; 肌肉無法被動的發育停止. 反而是, 非細胞間的心陳代謝作用, 其他週邊的組織和神經細胞被證明了:是自動連續性的神經而且帶來了合適的新陳代謝. 有很多的肌肉神經, 會在神經元內減少, (體細胞) 就是「色原溶解;核染質溶解;染色質溶解消失;色素溶解消失」也是代表了「官能障礙」
Perhaps the most striking evidence for the subtle but definite interdependence of peripheral structures and the central nervous system is provided by the stunted growth resulting from large cerebro-hemispheric and specifically parietal lobe lesions in the neonate human. This phenomenon was described by Macdonald Critchley (1955) and has been generally known to occur in connection with a condition called infantile hemiplegia. The stunting of the body side contralateral to the brain lesion occurs both in congenital and in acquired infantile hemiplegia and is not due to disuse, first because all tissues in the extremities involved are equally affected and second, because the arrest of growth starts at birth in the congenital cases, that is, before either the affected or the unaffected side is actively being used (Holt, 1961). These cases are even more interesting in the present context than the denervation atrohy, because here we see a relation between the highest level of the central nervous system and nonnervous structures of the periphery. Because the cells of the cerebral cortex are separated from those on the periphery by several internuncial neurons, there is an indication of a very subtle control that the higher centers of the central nervous system appear to exe3rcise upon the development of the body as a whole.
也許最驚人的事是:精細但與其他的結構相連,和由神奇的成長的中央神經系統
,在未滿月的)嬰兒時,會造成腦部的和顱頂骨腦的葉的器官損害。Macdonald 曾提過這種現象,而且他也想知道這種關聯:「未滿月的」嬰兒的未滿月的)嬰兒」。這種一連串的對於大腦的損害也會發生在先天的和後來得到的嬰兒的半身不遂,也不全是因為廢棄不使用的關係,而是因為末端的組織已經被感染。再者,在先天的生長案例中,還沒被感染的也會污染,這種狀況常常發生(Holt 1961)這狀況比起沒有脈落的萎縮現在的內容還有趣。因為我們看到了更高一樓的中央神經系統和其他的附近非神經的結構。因為大腦的皮質細胞是由不同的(神經細胞間之)聯絡的相連而成, 所以,更高一樓的中央神經系統是很精密的活動著,這也反映在整體的身體運作。
(b) Mechanical Relationships. In addition to the metabolic influences between nervous tissue and other tissue, morphogenesis is controlled by several other factors, some of which are indirectly related to neurophysiology. A good illustration is furnished by the mechanical forces exerted upon growing tissue, particularly bone, which stimulate cell division in certain directions. As muscles are innervated and begin to function, they exert a pull upon the boned to which they are attached and thus help shape the internal structure of this tissue.
(b) 機械學的,力學的;物理的關係
除了介於在神經組織和其他組織中的新陳代謝作用的影響之外,器官再生控制了幾個重要的因素,有些和神經生理學並沒有直接的關聯。作用暫成長的組織上的物理上力量,就是個好例子,特別是骨頭,在某種程度會刺激細胞分化。當肌肉該使受神經約束,該使運作時,會運作在附著的骨頭上,幫助使其形成內部組織。
It has often been noted (and, unfortunately, frequently been emphasized out of all proportion to its true importance) that the architecture of certain organs is ideally suited to their function. An excellent example of this was provided by D’Arcy Thompson (1942) who wrote, “In all the mechanical side of anatomy nothing can be more beautiful than the construction of a vulture’s metacarpal bone. The engineer sees in it a perfect Warren’s truss, just such a one as is often used for a main rib in an aeroplane.” The fundamental schema of the shape of individual bones and the skeleton as a whole are undoubtedly the result of evolutionary processes including selection and adaptation on a phylogenetic rather than an ontogenetic scale (Hackenbroch, 1957-1962). However, the actual realization of what is only potentially present in the fertilized egg is largely dependent upon factors which are active during ontogenesis. This is vest illustrated by the development of the internal structure of bones.
(很不幸的,常很少對於其重要性有成相對應的重視,有些器官的組成就是很符合本身的功能 )是很常被注意的。D’Arcy Thompson 就提供了個很好的例子,他寫道:『在解剖學的物理現象中,謀有什麼比禿鷹的手掌的骨頭更漂亮。這位工程師把它當成「養兔場的兔子」疝帶,托帶(Warren’s truss),就像飛機的主要肋骨架構』每跟骨頭的形狀的基本形狀都不同,而且,整個身體的支架也是進化的結果,包含了動植物種類史的選擇和突變,而不是個體發生的方向(Hachkenbroch,1957-1962)。但是,真正有可能在受精卵出現的時候,適一存在個體發生的過程中。骨頭的內部結構正好說明了這一點。
In 1866, a Swiss engineer, Culmann, noted that the internal trusses in the head of a human femur, anatomically known as trabeculae, were oriented in exactly the direction of the lines of maximum internal stress. He drew a diagram of a curved rod showing the lines of stress resulting from the application of a load from above (Fig. 1.1a). The model somewhat resembled the head of a derrick which he had just designed, and it is therefore referred to as a crane’s head. It bears a striking similarity to a section of the head of the femur (Fig. 1.1c). Culmann’s idea gave rise to J. Wolff’s famous theory expressed in his monograph on The Law of Bone Transformation (1870), according to which every change in the function of a bone produces changes in its trabecular architecture and external form in conformity with mathematical, static laws. The theory has since been criticized (Küntsher, 1934, 1936), modified (Murray, 1936), and elaborated upon (Evans, 1955, 1957, 1960; Carey, 1929). However, the idea is widely accepted today that muscles, through the tonus already present on early embryonic life, exert essential forces upon the growing bone. These pressures, together with those produced by differential growth of various parts of the embryo, result in stresses and strains which are the prerequisite stimuli for proper bone formation. [end]
在1866年,一個瑞士的工程師,Columann則對於人體的股骨的街頭內部分隔帶,則會在最大的內部壓力下,精確的呈現了其線條。他劃一個灣區的桿,則顯示壓力線下壓力會造成上面所提到的作用(Fig. 1.1a)。這種模式就好像他設計的油井的鐵支架一樣,也像個鶴的頭。它可以接受股骨的接頭部份所承受的壓力(Fig. 1.1c)。Culmann的想法影響了後來的J. Wolff有名的理論,顯現在他1870《骨頭的轉變法則》專題論文中 :在製造每個骨頭的功能時都會改變他的分隔帶的狀況,而且,外部形式時複合數學、統計法則的。(Kuntscher 1934,1936), 攻擊得理論,(Murry, 1936) 修改他,(Evans, 1955, 1957,1960; Carey, 1929)則把這理論發揚光大。但是,在今天大部人接受的肌肉理論,是藉由早期的初期/胎兒生物所呈現出肌肉強直性,在成長中的骨頭運作其基本的力量。這些壓力伴隨著胚胎不同階段的成長過程,會造成壓力,及對骨頭成長不可或缺的刺激的拉緊。
l New Words: Ontogeny: 【生】個體發生;個體發生學 Independent : 相互依賴的;互助的 ontogenetically: adj of ontogeny Ex-machina : is an American creator-owned comic book series created by Brian K. Vaughan and Tony Harris, and published by DC Comics under the Wildstorm imprint. Metabolic: 新陳代謝的 Trophic營養的;有關營養的 Regeneration:再生 contiguous連續的 limb 肢;臂;腳;翼; 大樹枝,主枝; 分支;執行者,代理人 vertebrates 脊椎動物 lizard: 蜥蜴 urodele salamander (傳說中生活在火中的)火蜥蜴,火蛇;火精,火怪 larval anuran, 幼蟲的;幼體的 anuran無尾目動物(蛙、蛤蟆等等) postmetamorphic :後變態的 amputated 醫】切斷;鋸掉;截(肢) stump口】腿;義肢[P] autograft 醫】自身移植物(如皮膚等) Morphogenesis, 器官發生 appendage, 生】附屬器官,附屬肢體(如臂、腿、尾等) mitotic 生】有絲分裂的 cartilage 【解】軟骨原骨,軟骨成骨 shriveled使束手無策;使無能為力;使無用 blastema 【生】胚芽 autonomic【生】自律的 embryology 胚胎學 denervation;去掉植】【動】脈序;脈狀;翅脈atrophy醫】萎縮;發育停止;虛脫 distal 【解】末梢部的;末端的 innervated使(器官、肌肉等)受神經支配 dystrophic (家庭或工業垃圾等對水的)污染 histological 組織學的 retrograde後退的;逆行的;退化的 soma【生】體細胞 chromatolysis【生】【醫】色原溶解;核染質溶解;染色質溶解消失;色素溶解消失 dysfunction. 官能不良;官能障礙
l cerebro-hemispheric 腦部的parietal【生】腔壁的;【解】顱頂骨的;【植】子房壁的 lobe【解】(腦、 肺等的)葉 lesions機能障礙;器官損害neonate (未滿月的)嬰兒 hemiplegia半身麻痺;半身不遂contralateral congenital天生的;先天的
l nternuncial 【解】(神經細胞間之)聯絡的 furnished 給(房間)配置(傢俱等);裝備[(+with)] metacarpal【解】掌的;掌部的 Warren’s truss Aeroplane【英】飛機[C] Femur解】股骨;【昆】股節 Trabeculae【解】分隔帶
l derrick:【機】轉臂起重機 / 油井的鐵架塔embryonic 胚芽的;胎兒的 prerequisite不可缺的;事先需要的;必修
LBT004-007惠珍 updated on Jan 5’ 09 (Done)
Chapter 1, section II)
II. FORM AND FUNCTION IN ONTOGENY
個體發生學的行是與功能
There is evidence (1) that the issues of the brain and the rest of the body constitute an organic, interdependent unit; and (2) that organisms are not programmed for their behavior by an ex-machina force, but instead they develop a program ontogenetically together with nervous and nonnervous tissues.
有證據顯示(1)「大腦的細胞組織」和「身體的其他的部分」是組成生物體這互相依賴的有機體。(2)這有機體並不是以他行為,或是「大機器」的力量(ex-machina force)來編碼的(programmed)> 而是,生物體根據神經和分神經組織而發展出「個體發生的編碼原則」。
(1) Mutual influence in the development of Nervous and Other Tissue
(1) 在神經和其他細胞相互影響的發展
Let us first consider the developmental relationship between nervous and other tissue. Our discussion can be divided into (a) metabolic or trophic relationships and (b) nonmetabolic, particularly mechanical relationhships.
讓我們把神經和其他細胞發展中玵互相影響的關係,考慮進去。我們的討論可分為2部。(a) 新陳代謝的或是營養的 (b)非新陳代謝的或是特定的物質上的關係。
(a) Trophic Relationships. Nervous tissue stands in an intimate relationship to other tissue anatomically contiguous to it. This is shown most clearly by the essential role played by nerves in the process of regeneration. There are a number of studies available which indicate that regeneration of an entire limb in lower vertebrates (fish, lizards, urodele, salamander, larval anuran, and postmetamorphic frog) and probably also in invertebrates is dependent on the presence of nerves in the amputated stump (Singer, 1959, Gutmann, 1964).
(a) 營養的關係。神經細胞和其他的很自然且連續在一起的細胞組織有很密切的關係。而這很明顯的顯示出在「再生」這過程,神經扮演了很基本的角色。有大量的文獻指出:「在低等脊椎動物上再生出一個肋骨」,無脊椎的如:(魚、蜥蜴、火蜥蜴、幼體無尾目動物、後變態的青蛙,及可能一些依賴神經存在的斷尾的部分。)
In a series of experiments by Schotté and Butler (1944), Singer (1947), and their students, and Nicholas (1949), it has been shown that an amputated limb will not regenerate unless an intact nerve is either present in the remaining stump from the beginning or is transplanted into the cut surface by autograft (however, see Thornton and Steen, 1962). Morphogenesis, that is, an orderly sequence of tissue differentiation and development of the lost appendage, will not ordinarily take place in the absence of living nervous issue during the very first stage of regeneration. If all nerves are removed form this stump during the earliest period, mitotic activity is dramatically slowed down; eventually some small amount of connective tissue, cartilage, and muscle may form in a disorderly nonfunctional fashion, giving a shriveled and shapeless appearance to the stump. If viable nervous tissue is not present from the start of the amputation but brought into the so-called blastema shortly afterward, regeneration takes place but the regenerate limb is poorly developed. The nerve need not be present throughout the entire period of regeneration; once the limb has begun to grow and tissues are sufficiently differentiated, the nerve may be removed without impariment of the morphegenetic potency acquired by these tissues during their earliest stage of formation. Singer (1974) has shown that it does not matter for regeneration what type of nerve, whether motor, sensory, or autonomic, is present in the blastema. It is merely the amount of nervous tissue present that controls the regenerative possibilities. Apparently a product of nerve-cell metabolism induces morphogenesis in the blastema.
在Scotte和 Bluter , Singer 和他們的學生及Nicholas,一系列的實驗中,顯示出:被截斷的肋骨是不會再生,除非他的神經一開始就仍然在存在的斷腿; 或是移植到自體移植皮膚表面的東西。(但,看看Thronton & Steen) 器官再生,就是細胞不同化和很多附屬器官的表層一連串的過程,在很早期的再生狀況下,缺乏活體神經組織之下,是不會發生的。如果所有的義肢、斷腿,都在早期都被移除,那麼有可絲分裂,就會大量下降活動; 甚至其他一些相關連的組織、軟骨、肌肉,都可能形成沒有規則、沒有功能的形狀、沒有用的、沒有形狀的斷腿的外觀。如果,神經細胞不是在開始截肢的時候就找到,是被帶到之後所謂的胚芽,「再生」就會開始。但是再生的肋骨市是「不完全發育」,神經不會在整個再生過程中呈現出來; 一旦,肋骨開始生長,而且組織得到足夠的養分,神經就可以從器官器官再生的部位移除,力量,是細胞在早期形成的時間得到的Singer,則證明了:任何細胞組織,對再生來說,是不相關的,不管是運動、感覺、自動的,都在胚芽中呈現。只是在細胞組織中呈現,然後控制再生的可能性,顯然地,細胞組織新陳代謝的解果會包括批牙器官再生。
These studies leave many questions about the biochemistry of embryology and growth unanswered; yet they do give us a glimpse of the complete interdependence and the natural integration of different tissues in the animal body. This impression is further strengthened if we consider some of the other trophic relationships that nerves have to peripheral tissue (for instance, the well-known fact of denervation atrophy). If the axon of a motor neuron is cut, the portion distal of the cut will die promptly, presumably because of its separation from its source of supply of vital substances (Gerard, 1950). But this is not the extent of the degenerative changes following the section of a motor nerve. The muscle innervated by the nerve will also undergo dystrophic changes with an extremely characteristic histological appearance. The loss of muscle substance is not due to a “functional” disturbance, such as the inhibition of nerve impulse transmission (Hamburger & Levi-Montalcini, 1950) nor due to disuse of the muscle; the muscle cannot be saved from atrophy by passive exercise. Indeed, the metabolic interdependence of nonnervous, peripheral tissue and nervous tissue is proven by the fact that the nerves themselves must have anatomic continuity with muscles for proper metabolic function. Severance of nerve from muscle will induce retrograde changes in the body of the neuron (the soma), known as chromatolysis, which is a sign of dysfunction.
對於生物化學胚胎學成長而言,對很多問題, 很多研究都還沒有有蒙卻的解答。但是, 這與給我們對於完全的依賴 或是再不同動物種類、細胞之間的整合演變,
有一種簡單的了解. 這也亙加強了我們對於其他的發育的關係的關感:細胞只是附屬的(例如:有名的去掉植)【動】脈序;脈狀;翅脈的例子). 如果運動神經的軸索被切斷, 末端的組織就會因為和濫以維生的物質分開而開始死亡。(Gerard1950) 但不是因為退化的物質, 改變運動神經. 而是受受神經支配的 肌肉會被大量的組織物質污染. 這種肌肉的損失是因為「功能亂流」例如;脈搏的跳動(Hamburger1950) 也不是因為肌肉組織造成的 ; 肌肉無法被動的發育停止. 反而是, 非細胞間的心陳代謝作用, 其他週邊的組織和神經細胞被證明了:是自動連續性的神經而且帶來了合適的新陳代謝. 有很多的肌肉神經, 會在神經元內減少, (體細胞) 就是「色原溶解;核染質溶解;染色質溶解消失;色素溶解消失」也是代表了「官能障礙」
Perhaps the most striking evidence for the subtle but definite interdependence of peripheral structures and the central nervous system is provided by the stunted growth resulting from large cerebro-hemispheric and specifically parietal lobe lesions in the neonate human. This phenomenon was described by Macdonald Critchley (1955) and has been generally known to occur in connection with a condition called infantile hemiplegia. The stunting of the body side contralateral to the brain lesion occurs both in congenital and in acquired infantile hemiplegia and is not due to disuse, first because all tissues in the extremities involved are equally affected and second, because the arrest of growth starts at birth in the congenital cases, that is, before either the affected or the unaffected side is actively being used (Holt, 1961). These cases are even more interesting in the present context than the denervation atrohy, because here we see a relation between the highest level of the central nervous system and nonnervous structures of the periphery. Because the cells of the cerebral cortex are separated from those on the periphery by several internuncial neurons, there is an indication of a very subtle control that the higher centers of the central nervous system appear to exe3rcise upon the development of the body as a whole.
也許最驚人的事是:精細但與其他的結構相連,和由神奇的成長的中央神經系統
,在未滿月的)嬰兒時,會造成腦部的和顱頂骨腦的葉的器官損害。Macdonald 曾提過這種現象,而且他也想知道這種關聯:「未滿月的」嬰兒的未滿月的)嬰兒」。這種一連串的對於大腦的損害也會發生在先天的和後來得到的嬰兒的半身不遂,也不全是因為廢棄不使用的關係,而是因為末端的組織已經被感染。再者,在先天的生長案例中,還沒被感染的也會污染,這種狀況常常發生(Holt 1961)這狀況比起沒有脈落的萎縮現在的內容還有趣。因為我們看到了更高一樓的中央神經系統和其他的附近非神經的結構。因為大腦的皮質細胞是由不同的(神經細胞間之)聯絡的相連而成, 所以,更高一樓的中央神經系統是很精密的活動著,這也反映在整體的身體運作。
(b) Mechanical Relationships. In addition to the metabolic influences between nervous tissue and other tissue, morphogenesis is controlled by several other factors, some of which are indirectly related to neurophysiology. A good illustration is furnished by the mechanical forces exerted upon growing tissue, particularly bone, which stimulate cell division in certain directions. As muscles are innervated and begin to function, they exert a pull upon the boned to which they are attached and thus help shape the internal structure of this tissue.
(b) 機械學的,力學的;物理的關係
除了介於在神經組織和其他組織中的新陳代謝作用的影響之外,器官再生控制了幾個重要的因素,有些和神經生理學並沒有直接的關聯。作用暫成長的組織上的物理上力量,就是個好例子,特別是骨頭,在某種程度會刺激細胞分化。當肌肉該使受神經約束,該使運作時,會運作在附著的骨頭上,幫助使其形成內部組織。
It has often been noted (and, unfortunately, frequently been emphasized out of all proportion to its true importance) that the architecture of certain organs is ideally suited to their function. An excellent example of this was provided by D’Arcy Thompson (1942) who wrote, “In all the mechanical side of anatomy nothing can be more beautiful than the construction of a vulture’s metacarpal bone. The engineer sees in it a perfect Warren’s truss, just such a one as is often used for a main rib in an aeroplane.” The fundamental schema of the shape of individual bones and the skeleton as a whole are undoubtedly the result of evolutionary processes including selection and adaptation on a phylogenetic rather than an ontogenetic scale (Hackenbroch, 1957-1962). However, the actual realization of what is only potentially present in the fertilized egg is largely dependent upon factors which are active during ontogenesis. This is vest illustrated by the development of the internal structure of bones.
(很不幸的,常很少對於其重要性有成相對應的重視,有些器官的組成就是很符合本身的功能 )是很常被注意的。D’Arcy Thompson 就提供了個很好的例子,他寫道:『在解剖學的物理現象中,謀有什麼比禿鷹的手掌的骨頭更漂亮。這位工程師把它當成「養兔場的兔子」疝帶,托帶(Warren’s truss),就像飛機的主要肋骨架構』每跟骨頭的形狀的基本形狀都不同,而且,整個身體的支架也是進化的結果,包含了動植物種類史的選擇和突變,而不是個體發生的方向(Hachkenbroch,1957-1962)。但是,真正有可能在受精卵出現的時候,適一存在個體發生的過程中。骨頭的內部結構正好說明了這一點。
In 1866, a Swiss engineer, Culmann, noted that the internal trusses in the head of a human femur, anatomically known as trabeculae, were oriented in exactly the direction of the lines of maximum internal stress. He drew a diagram of a curved rod showing the lines of stress resulting from the application of a load from above (Fig. 1.1a). The model somewhat resembled the head of a derrick which he had just designed, and it is therefore referred to as a crane’s head. It bears a striking similarity to a section of the head of the femur (Fig. 1.1c). Culmann’s idea gave rise to J. Wolff’s famous theory expressed in his monograph on The Law of Bone Transformation (1870), according to which every change in the function of a bone produces changes in its trabecular architecture and external form in conformity with mathematical, static laws. The theory has since been criticized (Küntsher, 1934, 1936), modified (Murray, 1936), and elaborated upon (Evans, 1955, 1957, 1960; Carey, 1929). However, the idea is widely accepted today that muscles, through the tonus already present on early embryonic life, exert essential forces upon the growing bone. These pressures, together with those produced by differential growth of various parts of the embryo, result in stresses and strains which are the prerequisite stimuli for proper bone formation. [end]
在1866年,一個瑞士的工程師,Columann則對於人體的股骨的街頭內部分隔帶,則會在最大的內部壓力下,精確的呈現了其線條。他劃一個灣區的桿,則顯示壓力線下壓力會造成上面所提到的作用(Fig. 1.1a)。這種模式就好像他設計的油井的鐵支架一樣,也像個鶴的頭。它可以接受股骨的接頭部份所承受的壓力(Fig. 1.1c)。Culmann的想法影響了後來的J. Wolff有名的理論,顯現在他1870《骨頭的轉變法則》專題論文中 :在製造每個骨頭的功能時都會改變他的分隔帶的狀況,而且,外部形式時複合數學、統計法則的。(Kuntscher 1934,1936), 攻擊得理論,(Murry, 1936) 修改他,(Evans, 1955, 1957,1960; Carey, 1929)則把這理論發揚光大。但是,在今天大部人接受的肌肉理論,是藉由早期的初期/胎兒生物所呈現出肌肉強直性,在成長中的骨頭運作其基本的力量。這些壓力伴隨著胚胎不同階段的成長過程,會造成壓力,及對骨頭成長不可或缺的刺激的拉緊。
l New Words: Ontogeny: 【生】個體發生;個體發生學 Independent : 相互依賴的;互助的 ontogenetically: adj of ontogeny Ex-machina : is an American creator-owned comic book series created by Brian K. Vaughan and Tony Harris, and published by DC Comics under the Wildstorm imprint. Metabolic: 新陳代謝的 Trophic營養的;有關營養的 Regeneration:再生 contiguous連續的 limb 肢;臂;腳;翼; 大樹枝,主枝; 分支;執行者,代理人 vertebrates 脊椎動物 lizard: 蜥蜴 urodele salamander (傳說中生活在火中的)火蜥蜴,火蛇;火精,火怪 larval anuran, 幼蟲的;幼體的 anuran無尾目動物(蛙、蛤蟆等等) postmetamorphic :後變態的 amputated 醫】切斷;鋸掉;截(肢) stump口】腿;義肢[P] autograft 醫】自身移植物(如皮膚等) Morphogenesis, 器官發生 appendage, 生】附屬器官,附屬肢體(如臂、腿、尾等) mitotic 生】有絲分裂的 cartilage 【解】軟骨原骨,軟骨成骨 shriveled使束手無策;使無能為力;使無用 blastema 【生】胚芽 autonomic【生】自律的 embryology 胚胎學 denervation;去掉植】【動】脈序;脈狀;翅脈atrophy醫】萎縮;發育停止;虛脫 distal 【解】末梢部的;末端的 innervated使(器官、肌肉等)受神經支配 dystrophic (家庭或工業垃圾等對水的)污染 histological 組織學的 retrograde後退的;逆行的;退化的 soma【生】體細胞 chromatolysis【生】【醫】色原溶解;核染質溶解;染色質溶解消失;色素溶解消失 dysfunction. 官能不良;官能障礙
l cerebro-hemispheric 腦部的parietal【生】腔壁的;【解】顱頂骨的;【植】子房壁的 lobe【解】(腦、 肺等的)葉 lesions機能障礙;器官損害neonate (未滿月的)嬰兒 hemiplegia半身麻痺;半身不遂contralateral congenital天生的;先天的
l nternuncial 【解】(神經細胞間之)聯絡的 furnished 給(房間)配置(傢俱等);裝備[(+with)] metacarpal【解】掌的;掌部的 Warren’s truss Aeroplane【英】飛機[C] Femur解】股骨;【昆】股節 Trabeculae【解】分隔帶
l derrick:【機】轉臂起重機 / 油井的鐵架塔embryonic 胚芽的;胎兒的 prerequisite不可缺的;事先需要的;必修
2008年12月2日 星期二
2008年12月1日 星期一
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