It is the influence of muscle tonus upon bone formation that brings this aspect of morphogenesis back to an interaction with the central nervous system. Muscle tonus is, in a sense, the beginning of behavior. Unless tonicity appears in a balanced, centrally regulated sequence, malformations of the outer and inner form of bones and limbs will result (as is indeed encountered in a number of well-known clinical conditions, for example, clubfoot deformities). Once more we see how bones, nerves, muscles, and behavior evolve as an organic entity. 就是肌肉強直性對骨骼形成的影響把形態發生的觀點拉回到與中央神經系統的互動中。肌肉強直性是、就某種意義來說、行為的開端。除非身體組織之正常官能出現在協調的、中央條理化的次序，不然骨骼與四肢外面與裡面形態的畸形就會產生（確實在多數眾所皆知臨床症狀中遇過、例如、畸形足）。我們再一次看到了骨骼、神經、肌肉、與行為是如何發展成有組織的實體。(2) The Embryology of Behavior Why should the metabolic and mechanical influences upon growth be of interest to the behaviorist? For all we know, this is an irrelevant detail of the constructional history of the organism that need not concern the student of behavior. We have had some indication, however, that some primitive precursor of behavior, particularly muscle tonus, is an important factor for at least one aspect of skeletal development. This raises the general question of the embryological history of behavior.(2) 行為的胚胎學 為何新陳代謝與物理的對生長的影響讓行為學家感興趣呢？就我們所知、這是不相關的並不會影響行為學者有機體建造由來的細節。然而，有些徵兆顯示一些行為的原始先驅，特別是肌肉強直性，至少是骨骼發展其中一方面的重要因素。這提出行為胚胎由來的普遍問題。 A complete survey of behavioral studies in prenatal development was prepared by Carmichael (1954) and should be consulted for details. In addition to observations on larval behavior in amphibians and pre-hatching behavior in chick embryos, an impressive amount of information exists on prenatal behavior in mammals, ranging from the primitive opossum to human fetuses. There is unanimity among observers that sensory irritability and motor activities come into play as soon as tissues have the differentiation and basic morphology which is characteristic of a given animal. In most mammals this tends to occur roughly toward the end of the first trimester of gestation. Much of the present-day thinking in this connection goes back to the pioneering work of Coghill (1929). 產前發展的行為研究之完整概論是由Carmichael所準備的且應該被當成細節的參考。除對兩棲動物的幼體行為及小孩胚胎孵化前的行為之觀察外，也有大量的令人印象深刻的哺乳類產前行為的資訊、從負鼠分佈到人類胎兒。觀察者皆一致同意對組織一有區分且有已知動物特色的基本的形態學，感覺的過敏性及運動的活動就會開始起作用。在大多數的哺乳類中，這大約傾向發生在接進懷孕期的前三個月底時。當今對此大多數的想法可以追溯至Coghill先導之研究。 Coghill observed the flexions and, later, the undulating movements in the trunk of the larval amlystoma, which are the behavioral primordium of the mature swimming movements. He demonstrated anatomically that the growth of the nervous system is directly correlated with the emergence of behavior, and he was able to explain certain features of the development of swimming movements (for example, that the movements of the limbs are at first strictly correlated with the gross movements of the trunk and only later assume a degree of independence) on strictly neuro-anatomic grounds. Coghill’s over-all conclusions were that behavior (in the animals studied by him and his students) emerged as an integrated but undifferentiated whole involving the entire animal from the start. With further development, a process of individuation of patterns appears which is manifested by the emergence of more specific and more detailed behavioral sequences; but these sequences were considered by him as parts of the behavioral schema as a whole which is potentially present from the beginning.
Coghill觀察了是幼蠑螈軀幹的彎曲部份及波動的活動（這是成熟游泳活動的行為原基）。他以解剖展示神經系統的發展與行為的出現是直接相關的，而且他在嚴格地神經解剖的基礎下能解釋某些游泳活動發展的特色（舉例來說，在一開始時，肢的活動跟軀幹的總活動是嚴格相關的，晚一點時才能取得獨立的程度）。Coghill全部的結論（他及他的學生所研究的動物）是以合併但是無差別的全體（從一開始時就牽涉了整個動物）出現。藉由更特殊的及更詳細的行為次序的出現讓個體化的樣本過程與進一步的發展顯露出來，但是這些次序他認為是行為概要（是從一開始就潛在呈現的全體）的一部份。 The concept of the integrated primordial whole, present from the beginning, has not been accepted by all embryologists (Kuo, 1939). Windle (1950) and his group, for instance, had reservations on this point; these workers tended to stress the developmental assembly of initially unrelated reflexes in to new and more complex patterns of coordination such as locomotion or feeding. When prenatal development of higher animals in considered, the controversy over the assembly view as against the differentiation view is quite academic. A survey of behavior embryology today leads to the following general conclusions.
從一開始就呈現的合併的原始全體這個概念並沒有被全部的胚胎學者接受（郭, 1939）。Windle（1950）及他的團體，舉例來說，對這點仍有保留；這些人員傾向強調從一開始不相關的反射作用發展裝配成新的及更複雜的協調樣本，像是移動或是餵養。當高等動物的產前發展被考慮時，反對分化觀點的裝配觀點是相當學術的。今日行為胚胎學的調查帶出了以下之一般結論。 First, the order in the emergence of sensory and motor capacities, reflexes, and spontaneous motoric events is constant and predictable for any given species. The sequence of events is not dependent upon experience.
第一，知覺的及運動能力、反射作用、及自發的運動事件之出現順序對任何特定之種類來說是不變及可預測的。事件的次序並不依賴經驗。 Second, the embryological emergence of behavior in mammals cannot be characterized by on simple scheme such as gradual differentiation of a whole or assembly of independently arising component reflexes. Although some aspects of behavior do emerge as undifferentiated patterns with subsequent individuation (for instance, the movements or limbs and digits and, to some extent, of head and trunk), other aspects of behavior are best regarded as a lawful but gradual integration of reflexes that can be observed to function independently before the total integration takes place. The natural history of gait in man is an example of the latter condition. Steeping movements and righting reflexes are present long before the infant is capable of coordinating them for competent stance or locomotion. Coronios (1933) described many similar examples in the course of his observations on the embryological development of behavior in the cat. But in addition to individuation and assembly, we may observe other types of reflex histories that fit neither of these two schemes. For instance, some motor reactions, such as the tonic neck reflex in man, emerge at a specific time only to disappear again in the course of further development. Other reflexes undergo certain transformations in function and appearance, such as infantile startle reactions, whereas still other types of reflexes may merely be suppressed by the appearance of inhibitory mechanisms, but can reappear if the inhibitory mechanisms are abolished; for instance, sucking and rooting reflexes may be seen in patients with advanced degenerative diseases affecting the cerebral cortex, in particular. The embryology of human behavior has been studied by Hooker (1952) and Humphrey (1964) who have also reviewed the recent literature on the subject. In man, certain synergisms, such as grasping, have been observed as early as the eleventh gestational week. After completion of the first trimester, that is , by the fourteenth week, most reflexes seem to be present that can also be observed in the neonate born at term.
第二，在胚胎學上，哺乳類行為的出現可以用一個簡單計劃來描述，像是一個全體的逐步分化或是獨立上升的反射作用的構成要素之裝配。雖然行為的一些方面的確是跟著隨後的個體化以分化的模式出現（舉例來說，肢及手指或足趾的移動而且，就某種程度來說，頭及驅幹的活動），其它行為的方面最好視為合法但是反射作用（觀察到可以在合併完成前獨立作用）的逐漸合併。人類步伐的自然歷史是後者條件的例子。步行移動及正確的反射作用在嬰兒為了充份地站立或運動的需要有能力去協調它們很久之前就出現了。在Coronois（1933）對貓的胚胎學行為發展觀察過程中，他描述了許多相似的例子。但是除了個體化及裝配外，我們能觀察其它無法符合這兩種模式的反射作用種類的歷史。舉例來說，有一些動作反應，像是人類在特定時間出現、只在進一步發展過程中再次消失的強直性頸反射。其它的反射作用在功能及外觀上經歷某種程度的轉化，像是嬰兒的驚嚇反應，而其它的反射也許僅僅被抑制機制的出現所壓制，但是如果抑制機制廢除，就能再出現；舉例來說，尤其可以在患有影響大腦皮層的高級退化疾病之病人中看到吸吮及尋乳反射。重新探討這個主題的文獻的Hooker (1952) 及Humphrey (1964) 研究了人類行為胚胎學。人類的某些連合作用，像是抓，最快在懷孕十一週時就會看到。在嬰兒出生到期可觀察到的反射，大部份的似乎在最先的三個月完成後、即十四週前就會呈現。 Third, species differ considerably in the order of emergence of spontaneous behavior. In most birds, for instance, the embryo may be seen to move before there is any sign of sensory reactivity. In mammals, however, sensory reactivity is very early, and in many species, including man, spontaneous behavior does not become prominent until after peripheral sensitivities have developed. There are, however, enough exceptions to this to discredit the notion that fetal behavior is the result of learning.
第三，物種在自然的舉止出現的順序上有相當大的不同。大部份的鳥，舉例來說，在有任何知覺反應的徵兆前就可以見到胚胎移動。然而，哺乳類的知覺反應是非常早地，且在大部份的物種中、包含人類，直到外周敏感度發展後，自然的舉止才變得比較重要。但是，除了這還是有足夠的例外去質疑胎兒的行為是學習的結果的這個概念。
The entire developmental process is of a physiological nature. In the earliest stages those metabolic events that have to do with cell division and tissue differentiation are most predominant. With the emergence of tissues differentiated into bone, muscle, and nerves, physiological interactions begin to manifest themselves as mechanical, motor events in addition to the biochemical events. Coordination emerges together with morphology, and a matrix for the animal’s future behavioral repertoire is developed, grows and develops into complex synergisms regardless of the “use” the individual will make of it in postnatal or mature life. This is not to imply that the habits of an individual at birth are so predetermined as to dominate completely his adult habits. Many modifications occur after birth in most vertebrates, but the scope of modifiability is always limited by genetic and prenatal events.這整個的發展過程有生理的本質。在最初的階段，那些跟細胞分裂及組織分化有關的新陳代謝的事件是最顯著的。隨著組織分化成骨骼、肌肉、及神經，生理的互動除了以生物化學的事件顯露之外，也開始以物理的及運動的事件顯露。協調與形態學一起出現，且不管個體在出生後或是成熟期如何使用，動物未來行為的全部功能之母體形成、成長及發展成複雜連合作用。這並不暗指個體出生時的習性如此預先確定以至於完全支配成年的習性。大部份的脊椎動物在出生後發生修改，但是修改的範圍總是受限於基因及產前的事件。
vocabulary
tonus (肌肉)強直性
morphogenis 1.形態發生 2.器官發生
tonicity 1.身體組織之正常官能 2.強壯 3.(肌肉)強直性
malformation 畸形; 殘廢
clubfoot 彎腳; 畸形足
deformity 畸形狀態
embryology 胚胎學
metabolic 變化的; 新陳代謝的
precusor 前導, 先驅, 前輩, 前兆
prenatal 產前的; 出生之前的, 胎兒期的
amphibian 兩棲動物
hatch 孵出
chick 小孩
opossum 負鼠
fetus 胎; (三個月後的)胎兒
unanimity 同意; 全體一致, 一致同意
trimester 三個月
gestation 懷孕; 懷孕期
undulating 波浪起伏的; 波浪形的
anatomically 解剖學上; 結構上
primordium 原基
locomotion 運動, 移動, 旅行
gait 步伐, 步態
infantile 幼稚的; 嬰兒的
degenerative 退化的; 變質的
synergism (各種藥物的)連合作用
neonate (未滿月的)嬰兒
fetal 胎兒的; 胎的
postnatal 產後的; 出生後的
vertebrate 脊椎動物
