LB 015-018 T

(3) Developmental History of the Central Regulatory Mechanism

The logical argument offered by Lashley is supported by an impressive array of experimental findings. We have mentioned the experiments on salamander larvae in which limb buds were transplanted to inappropriate sites. If a left forelimb is amputated from a donor animal and transplanted as a supernumerary limb to a host animal where it is allowed to regenerate into the right armpit, the extra limb is soon found to be moving smoothly. No tonic rigidity is noticed, and therefore we must assume that agonist and antagonist muscles receive innervation that is appropriate to the muscle. Interestingly enough, the limb will move at the tome that is appropriate for a forelimb to move; since, however, we have changed sides in the process of transplantation, the super numerary limb will move in the opposite direction from the original limb that is next to it. Thus one limb cancels the effect of the other, and it is possible to have a preparation with totally paradoxical behavior.

Lashley所提出的邏輯辯論是根據一個印象深刻的實驗結果.我們可以注意到在蠑螈幼蟲的實驗中,對於幼蟲四肢的解釋並不恰當.如果由捐贈器官的動物切除其左前肢,並移植到另一隻腳到宿主動物身上,則這隻額外移植的腳能馬上行動自如,沒有任何僵硬或不適的情形,因此,我們可以假設收縮筋和頡頏肌的神經分布是適合肌肉生長的.有趣的是,前肢移動的時候也會帶動後肢的移動,然而,當我們將移植改變到另一側時,額外移植的另一隻腳則會往反方向移動.因此,一隻腳相對抵消了對另外一隻腳的影響.這對動物不合常理的行為提出了一種可能的解釋.

What is the nature of this relationship between the limb and the brain? How can reciprocal innervation of muscles and timing of the limb with respect to other limbs be established in a fairly orderly way where there could not have been any neuronal “wiring” for the additional leg? Inspection under the microscope of the regenerated tissues does not reveal any visible order. Never fibers seem to have sprouted every which way, and the established connections seem to be entirely random. Could this be a delusion due perhaps to insufficient power of resolution of the light microscope? Is it possible that the nerve sprouts actually find their way to the appropriate muscle because of some unknown biochemical affinity between muscle and nerve? At first this possibility was never entertained. Instead it was thought that muscles were physiologically tuned to specific neuronal messages and simply responded whenever they “heard their name over the public address system.” This hypothesis was known as the muscle-resonance theory. However, Wiersma(1931) disproved the theory by recording electrical potentials from the nerves. Subsequently, the orderly recovery of motor coordination in the transplanted limb was interpreted on the basis of structural connections. There are two essential possibilities here. Either the nervous system entirely fixed and proper connections are made at the periphery in the way first mentioned, that is, fibers that carry given messages have the capacity of finding their way into the appropriate muscle during regeneration; or the muscles have the capacity of influencing the nerves that grow into them and thus affect the central nervous system retrogradely.

而四肢和腦部之間在本質上存在著什麼樣的關係呢?肌肉組織的神經交互分布和四肢之間的關聯性要如何建立一個相當有秩序的方式,讓那隻額外移植的腳沒有發上任何神經系統上錯誤的?把肌肉重生的組織放在顯微鏡下觀察,似乎無法從肉眼得知.神經組織可能以任何方式生長以及發展,而其中所建立的連結似乎是隨機的.這會是由於顯微鏡的觀察沒有提供充分的解答所造成的誤解嗎?神經系統的生長有沒有可能是因為肌肉組織和神經之間存在生物化學上,異種物質間起化學作用所產生的吸引力?起初,這種可能性是完全不被接受的.當時是認為肌肉在生理學上調整成特有的神經訊息,只單純地在公開的演講系統聽到他們的命名時作回應,這樣的說法就是著名的肌肉共振理論.然而,Wierma(1931)以神經的電位來推論此理論.其次,針對經過一值得繳,其運動神經協調性的復原,說明了神經連結架構的基礎.在這裡,有兩種重要的可能性.第一種,神經系統是固有的,而其中的連結是根據一開始所提到的外圍方式,也就是說,在肌肉重生的過程中,帶有訊息的組織有能力去尋找到適合的肌肉組織.另外一種可能性是,肌肉擁有影響神經的能力,藉以降低中樞神經系統的影響力.

The first of these two possibilities has gained plausibility in most recent investigations (Mark, 1965) , although it is still far from established. The second possibility is favored by many of the neuroembryologists who had made the original discoveries on lower vertebrates. In Weiss’s own words (1950b) : It is thought now that “each muscle has a specific biochemical differential, that it projects this differential into the motor nerve fibers that come to innervate it and thus tunes (modulates) the motor ganglion cells to a specificity appropriate for the particular muscle. The ganglion cells have received their specificity by a retrograde influence (modulation) from the muscle itself.” Until recently, Sperry (1958) believed that the biochemical influence exerted by the muscle upon the nerve actually induces synaptic changes in the central nervous system. But Eccles et al. (1962) found only limited support for this interpretation, lending credence to Mark’s (1965) interpretation, a point of view that is also now favored by Sperry (1963). For an up-to-date review of the entire topic see Weiss (1965).
此兩種假設雖然和以建立的理論有一段差距,但仍在之後的研究中逐漸取得合理性.其中,第二種假設最受到許多從事研究較低階脊椎動物的神經病理學者支持.在Weiss所提出的論述中:每一條肌肉都具有生物化學上的差異,並將這樣的差異性投射於運動神經的組織中,並且改變運動神經節細胞,成為只針對某特定肌肉發展的細胞.神經節細胞接收到這樣的訊息是受到肌肉本身調整的影響.一直到西元1958年,Sperry相信運用神經肌肉在生物化學上的影響,在中樞神經系統上確實減少了生殖細胞在進行減數分裂的前期,相同染色體互相結合的現象.但是,Eccles et.al在西元1962年發現只有少數研究支持這項理論,而對於Mark在西元1965年的研究給予肯定.同時也獲得了Sperry的支持.相關最新的且完整的論述可以參考Weiss (1965).

The importance of the original discovery is that in phylogenetically primitive vertebrates (and probably during fetal stages of most other vertebrates) there is an inescapable BaupIan (blueprint) for both the gross form and the sensory-motor integration. The surgical rearrangement experiments on lower forms show how difficult it is to interfere with the “preestablished harmony” of the movements of muscles throughout the body which accounts for smooth coordination.

最初的重要發現是對於脊椎動物一開始有關物種的變化(或者是在大多數脊椎動物的胚胎階段)有一個關於數量以及運動肌感知的整合性的藍圖.在較低階的外科手術重组實驗中顯示在流暢的身體協調度上肌肉運動前期在協調方面的困難

Compare this situation with rearrangement experiments in mammals and adult forms of lower vertebrates. If the nerves which normally feed a flexor and extensor pair of muscles, respectively, are interchanged surgically and are allowed to regenerate into the wrong muscle, subsequent coordination becomes disordered and remains so.

和較低階的脊椎動物中的哺乳類動物重组實驗的相比較之下,如果屈肌和伸肌在外科手術上互相交換,並且重新生長錯誤的肌肉,則協調性會受到阻礙,且阻礙的形況會一直持續下去.

The difference in the results of rearrangement between lower and higher forms is not as paradoxical as it might appear at first. Table 1.1 summarizes the situation for easier reference. We discern here the emergence of a specific theme. For all animals examined, rigid plans for development of form and motor coordination seem to exist. In primitive forms, tissues are less differentiated or specialized and thus participate in the organization responsible for motor coordination; end organs may influence the structure and function of centers as much as the centers may influence the periphery. The result is preservation of the original plan for integration. In adult and higher forms, tissues become more and more specialized and thus more independent of each other. The motor-integration plan is no longer “inscribed” in tissues other than those directly concerned with coordination, principally the brain. The basic plan or plans (the dispatch schedules) for sensory motor coordination are still as rigidly inherent in the internal organization of the animal but they are stored now in the central nervous system alone. In this context, the dimension of plasticity-rigidity refers exclusively to adaptation and readjustment of internal process, not to an animal’s adaptation to environmental conditions.
在較高階和低階的層次上,重组結果的不同,並非是不合理的.圖表1.1說明了先前所參考的情況.我們認知到了特殊命題的急迫性.在所有調查的動物中,較堅硬植物的生長和運動神經的協調性是存在的.對於運動神經的協調度而言,組織初期的形狀並無明顯不同.而最終器官或許會影響組織核心的架構和功能,就像是組織核心影響外圍一樣.其結果是在成熟或更高階的形式,組織變得越來越有個別性,越來越獨立於其他組織.而運動神經整合的計畫不再只是存在組織當中,而是直接參與協調.主要以腦部為主.最初對於感知運動在神經協調性的計畫,是嚴格地經由遺傳到動物的內部組織中.但它們現在仍獨立儲存於中樞神經系統中.在這裡,適應性和剛性所指的是內部過程中的適應性和重新調整的能力,而不是指動物對外界環境的適應性.

LB 453-455 T

John Locke (1632-1704) discussed language because he had found that,” There is so close a connection between ideas and words…that it is impossible to speak clearly and distinctly of our knowledge, which all consists of propositions, without considering first, the nature, use, and signification of language “ which he did not equate with reason. God had” designed man for a sociable creature, ...under a necessity to have fellowship with those of his own kind;…furnished him also with language…the greatest instrument, and common tie of society.” To that end man’s organs were fashioned “ to form articulate sounds “and he was given the ability “to use these sounds as signs of internal conceptions, and to make them stand as marks for the ideas within his own mind…” Reading Locke up to this point, one may have gained the impression that he regarded language and society as creations of God. However, he continues in the next chapter:

John Locke 討論到語言,因為他發現思想和語言之間有著密不可分的關係.不可能和我們的知識清楚地區隔開來.因為我們的知識包含了語言的本質,使用方式和文字本身的意義.在此,他並沒有將理性加入討論.上帝將人創造成具社會性的受造物,同時提供人類語言,因為語言是最好的工具,也是共同的社會約束.為了讓語言成為共同的社會約束,人類的發音器官被上帝塑造成是用來產生發音清晰的聲音,並同時賦予人類用聲音表達內在思想的能力,讓聲音成為人們表達思想的符號. 在看完Locke的論述之後,或許有人認為他將語言和社會視為上帝的創造物,然而,他在下一個章節繼續談論到:

“The comfort and advantage of society, not being to be had without communication of thoughts, it war necessary that man should find out some external sensible signs, whereby those invisible ideas… might be known to others. For this purpose, nothing was so fit, either for plenty or quickness, as those articulate sounds, which, with so much ease and variety, he found himself able to make.”[48]

「源自社會的安適和優點,是起因於思想的溝通,人類必須要藉由無形的或者是他人的思想察覺到來自外界可感知的符號,為了達到思想溝通的目的,發音清晰的聲音是再適合不過的了.因為人類可以輕而易舉地發出聲音,並且產生不同的變化.」

So that language had been “found” by man in order to benefit from “the comfort and advantages of society” and no longer had he been “designed as a social creature” and simply been “furnished with language.” Locke appears to have abandoned the conviction of the importance of the social factor in the origin of language when he later discussed the invention of words by Adam. (According to the Bible Adam was alone when he named the animals) [49]. Locke has been extensively quoted, because the widely held thesis that he saw the origin of language as a free act of invention[50] does not appear acceptable without some reservations.¹


所以,人類「發明」語言是為了要能夠得益於社會的安適和優點,人類不再只是上帝加上語言後所創造出來的受造物.從語言起源的角度來看,在Locke談論到Adam發明文字的過程,他被認為是放棄了「社會」這個重要因素.(因為根據聖經的說法,Adam是獨自幫動物命名的)而Locke甚至更擴大引用當時所廣泛支持的論點,語言的起源是自然而然產生的創造行為.


A defender of Locke’s sensualism, the Abbé Etiénne Bonnot de Condillac (1715-1780) thought that man’s first language consisted of gestures and inarticulated noises which were“ based on the construction of our bodily instruments”[51].This gesture language already contained artificially chosen and used signs. But Condillac is careful to differentiate artificially created from willfully chosen signs.² Articulated sounds were used initially to emphasize gesture language. Then sounds came to be used in imitation of natural sounds. Gradually, as articulated sounds were increasingly used, gesture language was replaced by articulated language. This transition was favored by the discovery that sounds are suitable for the expression of the physical characteristics of objects. The first names did not contain any truth about object, for they reflected only peoples’ impressions of objects, not the nature of the object itself [52]. The understanding of gesture language, like the comprehension of a picture had required the application of analysis and the use of analogy. Articulated language differed from gesture language only in one respect. Ideas were presented in succession instead of simultaneously [53].³

孔狄亞克,Locke感覺論的擁護者,他認為人類最初的語言包含了以人體結構為基礎的手勢以及含糊,發音不清楚的聲音.這種含有手勢的語言已經包括了人為的選擇和習慣性.在此,孔狄亞克對於人為的創造以及具有目的性所選擇的聲音有另外再加以區別.最初人類會發出聲音是為了要強調動作.接著,聲音開始使用於對自然的聲音的模仿.逐漸地,隨著聲音的使用頻率增加,手語便被聲音所取代.這樣的轉變,最主要是因為發現到以人體結構的特徵使用聲音更為適合.一個物體最初的名稱並不包含其本質,只是人類對其物體的印象的一種反映.對於手語的理解,就像人類對於物體的印象,必須應用分析和類比的概念.使用含有語音的語言和手語唯一的不同點在於聲音是連續的,而不是同時間發生的.

Although Nature gave man “nearly complete freedom to do as he wills” in articulated language, she” guides us by putting the first sounds in our mouths and we discover other sounds by analogy” [54].Condillac concluded by saying:

雖然大自然給予人類在發音時,以其自由意識為依歸,近乎完全的自由.祂將最初的聲音放入人類的嘴中,並讓我們用類比的方式發現其他的聲音,孔狄亞克作了結論:

“I have said enough… to show that languages are the work of nature, that they were formed so to say, without us and that we, as we worked at language, blindly followed our way of seeing and feeling.”
Language is for man the result of an inner need, just as it is in children. Once we begun to speak, we have to drive to enrich our language with new expressions [55]. The inquiry into the basic principles of language would have to consider first that language which we have by virtue of our “bodily organization.” Once we shall have discovered the principles according to which we speak, we shall understand the rules for speaking any other language [56]. The organic nature of language or its biological basis was apparently not just a theoretical concept to Condillac, for he saw the need to investigate and apply it to the study of languages.

語言對於人類而言,是呈現內在需求的結果,就像在孩童時期一樣,當我們開始學習說話,我們就被迫使用各種不同的說話方式.在探討語言的基本要素時,首先第一點必須要考慮到的是,以我們身體器官的優點來討論語言,在之前曾經討論過我們在講話時的原理,就應該了解到,在說各種不同語言時的規則,不論是從語言的起源或者是生物基礎的角度來看,孔狄亞克的論述是不夠的,因為他只看到了研究語言的需要,並且將其應用於語言習得上.

In the opinion of George Louis Leclerc de Buffon (1707-1788), a specifically active sense of hearing in man plays an important role in language formation. Articulation is considered less important, in view of the belief that apes also have organs for articulation. All people speak naturally, and Buffon is the first to make this the primary difference between man and animal. Man spoke because he had reason. Animals do not have language because they lack man’s ability to think and cannot connect concepts [57].
Even for the author of the “Nature History,” reason seemed to be the most basic aspect of language. However, in the mid-eighteenth century man’s reason was, strictly speaking, not considered a part of nature. A more naturalistic approach is expressed in the “Lectures on the Theory of Language and Universal Grammar,” publisher in 1762 by Joseph Priestley (1733-1804). All social creatures have a God-given way of communication, and languages are like plants, which grow, blossom, and then wilt. The complexity of a language is never the result of design, but is due to accident and the structure of Man’s speech organs. Languages are subject to natural law ,therefore one should not attempt to fix strict rules for its usage [58].

在Buffon的觀點來看,特別是人類主動的聽覺能力在建構語言上扮演著非常重要的角色,清晰的發音則是其次.因為,一般認為,猿猴也有發音器官,而所有人類都能自然地說話,Buffon是第一位將其視為人類和動物最大的差異.人會說話是因為擁有理性,而動物沒有語言是因為牠們缺乏像人類一樣的思考以及溝通能力.
即使是《Natural History》的作者,Buffon,亦將理性視為語言當中最基礎的一部分.然而,在18世紀中葉,人類的理性,更嚴格地說,不再被視為本質的一部分,在1762年,Joseph Priestley所出版的”Lectures on the Theory of Language and Universal Grammar”「語言理論.普遍語法」中,自然主義可見一斑.人類擁有上帝所賦予的溝通的天賦,而語言就像植物一樣,會成長,茁壯,凋零.語言之所以會複雜是由於人類發音器官的內部結構.語言是屬於自然法則的一部分,任何人都不應該企圖用嚴格的規則去規範它的用法.