Pathways in the emergence of developmental neuroethology: Antecedents to current views of neurobehavioral ontogeny

The historical forces that have contributed to our current views of neurobehavioral development (and thus to the fields of developmental psychobiology and neuroethology) are many and varied. Although similar statements might be made about almost any field of science, it is in particular true of this field, which represents a kind of mongrel discipline derived from at least three major sources (psychology, embryology, and neuroscience) and several more minor ones (including developmental psychology and psychiatry, psychoanalysis, education, zoology, ethology, and sociology). Although I attempt to demonstrate here how each of these sources may have influenced the emergence of a unified field of developmental psychobiology or developmental neuroethology, because the present article represents the first attempt of which I am aware to trace the history of these fields I am certain that there is considerable room for improvement, correction, and revision of the views expressed here. Accordingly, I consider this inaugural effort a kind of reconnaissance intended to trace a necessarily imperfect historic path for others to follow and improve upon. In the final analysis, I will be satisfied if this article only serves to underscore two related points: first is the value derived from historical studies of contemporary issues in development, and the second concerns the extent to which our current ideas and concepts about neurobehavioral development, ideas often considered new and contemporary, were already well known to those who came before us. The first point underscores the arguments expressed in the Introduction that the present must always be reconciled with the past, for the past is never entirely past. The second point returns full circle to an important thought expressed in the opening quotation to this article, namely, that even though our historic predecessors lacked much of the empirical facts available to us they were nonetheless able to attain a surprisingly deep understanding of neurobehavioral ontogeny. © 1992 John Wiley & Sons, Inc.     

Development of fluctuating asymmetry in tail feathers of the barn swallow Hirundo rustica

Fluctuating asymmetry represents usually small, random deviations from symmetry in bilateral morphological characters. The ontogeny of asymmetry in morphological characters may reveal information about developmental processes in a general sense. I studied the development of fluctuating asymmetry in feather characters of the barn swallow Hirundo rustica, that are developed repeatedly during the single annual moult, with the following results. First, the side developing a larger feather was found to be partially biased, as demonstrated by one side consistently developing a larger feather under natural and experimentally induced growth episode events. Second, asymmetric feathers were found to consist of asymmetric daily growth increments, and the size of the increments developing under different environmental conditions were positively correlated. Third, fluctuating asymmetries of feathers developing under different environmental conditions were positively correlated, although the level of asymmetry was larger under adverse environmental conditions. Fourth, individual asymmetries in tail length and growth bar length were unrelated to the duration of the developmental period, although late growth increments were smaller and more symmetric than early increments. These observations suggest that fluctuating asymmetry partially arises as a consequence of a random bias in the feather follicles and differences in environmental conditions during ontogeny of feathers.     

A simple model of the relationship between asymmetry and developmental stability

The relationship between developmental stability and morphological asymmetry is derived under the standard view that structures on each side of an individual develop independently and are normally distributed. I use developmental variance of sizes of parts, V_D, as the converse of developmental stability, and assume that V_D follows a gamma distribution. Repeatability of asymmetry, a measure of how informative asymmetry is about V_D, is quite insensitive to the variance in V_D, for example only reaching 20% when the coefficient of variation of V_D is 100%. The coefficient of variation of asymmetry, CV_FA, also increases very slowly with increasing population variation in V_D. CV_FA values from empirical data are sometimes over 100%, implying that developmental stability is sometimes more variable than any previously studied type of trait. This result suggests that alternatives to this model may be needed.     

The beak of the other finch: coevolution of genetic covariance structure and developmental modularity during adaptive evolution

The link between adaptation and evolutionary change remains the most central and least understood evolutionary problem. Rapid evolution and diversification of avian beaks is a textbook example of such a link, yet the mechanisms that enable beak''s precise adaptation and extensive adaptability are poorly understood. Often observed rapid evolutionary change in beaks is particularly puzzling in light of the neo-Darwinian model that necessitates coordinated changes in developmentally distinct precursors and correspondence between functional and genetic modularity, which should preclude evolutionary diversification. I show that during first 19 generations after colonization of a novel environment, house finches (Carpodacus mexicanus) express an array of distinct, but adaptively equivalent beak morphologies—a result of compensatory developmental interactions between beak length and width in accommodating microevolutionary change in beak depth. Directional selection was largely confined to the elimination of extremes formed by these developmental interactions, while long-term stabilizing selection along a single axis—beak depth—was mirrored in the structure of beak''s additive genetic covariance. These results emphasize three principal points. First, additive genetic covariance structure may represent a historical record of the most recurrent developmental and functional interactions. Second, adaptive equivalence of beak configurations shields genetic and developmental variation in individual components from depletion by natural selection. Third, compensatory developmental interactions among beak components can generate rapid reorganization of beak morphology under novel conditions and thus greatly facilitate both the evolution of precise adaptation and extensive diversification, thereby linking adaptation and adaptability in this classic example of Darwinian evolution.     

Effects of the egg incubation environment on turtle carapace development

Developing organisms are often exposed to fluctuating environments that destabilize tissue-scale processes and induce abnormal phenotypes. This might be common in species that lay eggs in the external environment and with little parental care, such as many reptiles. In turtles, morphological development has provided striking examples of abnormal phenotypic patterns, though the influence of the environment remains unclear. To this end, we compared fluctuating asymmetry, as a proxy for developmental instability, in turtle hatchlings incubated in controlled laboratory and unstable natural conditions. Wild and laboratory hatchlings featured similar proportions of supernumerary scales (scutes) on the dorsal shell (carapace). Such abnormal scutes likely elevated shape asymmetry, which was highest in natural nests. Moreover, we tested the hypothesis that hot and dry environments cause abnormal scute formation by subjecting eggs to a range of hydric and thermal laboratory incubation regimes. Shape asymmetry was similar in hatchlings incubated at five constant temperatures (26–30°C). A hot (30°C) and severely Dry substrate yielded smaller hatchlings but scutes were not overtly affected. Our study suggests that changing nest environments contribute to fluctuating asymmetry in egg-laying reptiles, while clarifying the conditions at which turtle shell development remains buffered from the external environment.     

Linear and threshold-dependent expression of secondary sexual traits in the same individual: insights from a horned beetle (Coleoptera: Scarabaeidae)

Elaborate horns or horn‐like structures in male scarab beetles commonly scale with body size either (a) in a linear fashion with horn size increasing relatively faster than body size or (b) in a threshold‐dependent, sigmoid fashion; that is, males smaller than a certain critical body size develop no or only rudimentary horns, whereas males larger than the threshold size express fully developed horns. The development of linear vs. sigmoid scaling relationships is thought to require fundamentally different regulatory mechanisms. Here we show that such disparate regulatory mechanisms may co‐occur in the same individual. Large males of the south‐east Asian Onthophagus (Proagoderus) watanabei (Ochi & Kon) (Scarabaeidae, Onthophagini) develop a pair of long, curved head horns as well as a single thoracic horn. We show that unlike paired head horns in a large number of Onthophagus species, in O. watanabei the relationship between head horns and body size is best explained by a linear model. Large males develop disproportionately longer horns than small males, but the difference in relative horn sizes across the range of body sizes is small compared to other Onthophagus species. However, the scaling relationship between the thoracic horn and body size is best explained by a strongly sigmoid model. Only males above a certain body size threshold express a thoracic horn and males smaller than this threshold express no horn at all. We found a significant positive correlation between head and thoracic horn length residuals, contrary to what would be expected if a resource allocation tradeoff during larval development would influence the length of both horn types. Our results suggest that the scaling relationship between body size and horn length, and the developmental regulation underlying these scaling relationships, may be quite different for different horns, even though these horns may develop in the same individual. We discuss our results in the context of the developmental biology of secondary sexual traits in beetles. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 473–480.     

赤眼蜂发育速率对梯度恒温的响应

赤眼蜂发育速率对梯度恒温的响应关系到生物防治效果问题,以稻螟赤眼蜂与玉米螟赤眼蜂为例,研究了两种赤眼蜂分别在10、15、20、25、30、35、37℃梯度恒温下的相应世代发育历期,并运用直线回归模型与王-兰-丁模型两种拟合法,对温度与发育速率的关系进行了拟合。研究结果表明:温度对两种赤眼蜂的世代发育历期有明显的影响,在试验梯度温度10-37℃范围内,两种赤眼蜂的世代发育历期随温度升高而逐渐缩短。在10℃至30℃范围内,两种赤眼蜂的发育历期迅速缩短。在30℃到37℃范围内,两种赤眼蜂的发育历期变化进入一个相对平稳时期。在试验温度为30、35、37℃,稻螟赤眼蜂与玉米螟赤眼蜂的发育历期分别为7.75、7.45、7.95d和8.05、7.50、7.90d,差异不显著。通过直线回归法,可得稻螟赤眼蜂与玉米螟赤眼蜂的发育起点温度分别为5.23℃和5.24℃,有效积温分别为212.77d·℃和217.39d·℃;利用王氏模型法,得到两种赤眼蜂的发育最低温、发育最适温和发育最高温分别为7.96、18.91、38.99℃和6.97、18.87、38.99℃。稻螟赤眼蜂与玉米螟赤眼蜂在10-37℃的试验温度范围内,发育速率随温度升高而相应增大。在10-30℃范围内,两种赤眼蜂的发育速率迅速增长;在30-35℃范围内,两种赤眼蜂的发育速率进入一个相对平稳时期,超过临界高温TH后,随温度升高发育速率最终降为0。讨论了赤眼蜂发育速率对梯度恒温响应研究中应注意的梯度恒温设计和数学模型选择问题。     

小黑瓢虫新猎物--高氏瘤粉虱发育速率模型的研究

高氏瘤粉虱Aleurotuberculatus takahashi David et Subramaniam是小黑瓢虫Delphastus catalinae的新猎物寄主,本测定了不同温度下高氏瘤粉虱的发育速率,得出高氏瘤粉虱各虫态的发育起点温度为：卵期8.30℃、一龄若虫6.46℃、二龄若虫7.71℃、三龄若虫7.12℃、四龄若虫一拟蛹7.68℃、产卵前期7.90℃、世代7.64℃。另外还得出发育速率与温度关系的线性日度模型、Logistic模型。     

Historical and philosophical perspectives on the study of developmental bias

Throughout the recent history of research at the intersection of evolution and development, notions such as developmental constraint, evolutionary novelty, and evolvability have been prominent, but the term “developmental bias” has scarcely been used. And one may even doubt whether a unique and principled definition of bias is possible. I argue that the concept of developmental bias can still play a vital scientific role by means of setting an explanatory agenda that motivates investigation and guides the formulation of integrative explanatory frameworks. Less crucial is a definition that would classify patterns of phenotypic variation and unify variational patterns involving different traits and taxa as all being “bias.” Instead, what we should want is a concept that generates intellectual identity across various researchers, and that unites the diverse fields and approaches relevant to the study of developmental bias, from paleontology to behavioral biology. I point to some advantages of conducting research specifically under the label of “developmental bias,” compared with employing other, more common terms such as “evolvability.”     

Commentary on the Role of Maternal Toxicity on Developmental Toxicity

Maternal mammalian toxicity impacts prenatal development, with general systemic maternal toxicity, from reduced weight gain to morbidity, causative for reduced fetal weights/litter and increased fetal variations (especially skeletal)/litter, but not, in the author's opinion, for increased fetal malformations, reduced litter sizes or full litter losses. Increased fetal malformations are likely due to exposure to specific chemicals which alter specific maternal functions at critical point(s) in pregnancy, typically exaggerated effects from higher doses by drugs under development with known, desired pharmacological effects. Malformations can also be from genetic/epigenetic alterations, specific altered proteins, molecular pathways, etc. Full litter losses are triggered by the mother and are rare in rats. Information to inform maternal (and developmental) toxicity includes ovarian corpora lutea counts, uterine implantation profile, degree of litter reduction (if present), timing and extent of maternal toxicity relative to those of adverse embryofetal effects, etc. The view of maternal toxicity as confounding results in in vivo developmental toxicity studies, worldwide concerns about increased research animal usage, increasing time, labor, costs, and new software and hardware sophistication all drive the interest in development, validation, and performance of in vitro/in silico assays. These assays are fast, inexpensive, responsive to animal use concerns and amenable to mechanistic questions. The strength of these in vitro/in silico assays is considered by many to be the absence of the maternal organism/placenta. These assays inform mechanism and hazard, but NOT risk. The Environmental Protection Agency currently estimates that these new assays are approximately 70% accurate versus the whole animal tests.     

Neurotoxicological effects of deltamethrin on the postnatal development of cerebellum of rat

Deitamethrin (DLT) has been accepted to be 10,000 times less toxic to man than to insects. While toxicity of DLT in adult animals has been studied using biochemical and electrophysiological tools, reports on its developmental neurotoxicity are rather scanty. Wistar rat pups were exposed to DLT (0·7 mg/kg body wt/day, i.p., dissolved in propylene glycol from postnatal day 9–13. Equal number of age matched pups were used as vehicle controls. The animals were weighed and perfused intracardially on postnatal days 12,15,21, and 30 and their brains dissected out. Cerebellum along with the brainstem was separated by a transverse section at the tectal level and processed for morphometric and toxicological studies. The micro-and inter-neurons in the cerebellum are known to differentiate and mature, both morphologically and biochemically, during the postnatal life of rats. Postnatal exposure to DLT has been observed to delay the cytogenesis and morphogenesis of these neurons. In addition to this, damage to the developing vasculature has also been recorded in the form of thrombus and haemorrhage. Focal degeneration and spongy appearance of the tissue in the vicinity of the damaged blood vessels have also been recorded. The study has opened up several questions on the safety of this substance to the pregnant mothers and infants in the habitats where this substance is in use for vector control.     

Relationships among development, ecology, and morphology in the evolution of Echinoderm larvae and life cycles

The evolution of life cycles involves transitions between discrete states in one or more of the characters that comprise a developmental pattern. In this paper, we examine three of the major life cycle characters and the states for these characters. Using examples from echinoderms, we discuss the evolutionary transitions that have occurred in the type of morphogenesis, developmental habitat, and mode of nutrition during development. We evaluate the functional requirements associated with these transitions to infer the likelihood (frequency or rapidity) of change in a given character and of biases in the polarity of character state transitions. Using comparisons of closely related species, we evaluate the change between states in one character for dependence on the state of, or correlated changes in, other characters. Based on our analysis of congeneric species that differ in developmental habitat, we conclude that the transition between pelagic and benthic development is an ecological change that is independent of changes in morphogenesis and should be reversible. In contrast, the transition from feeding to nonfeeding development has been considered to be irreversible because it involves marked changes in larval morphology. We re-examine the transition between different modes of larval nutrition in light of recent studies that show that there exists a continuum of nutritional strategies between planktotrophy and lecithotrophy. This continuum is largely determined by variation in maternal investment and does not involve alterations in larval morphology. We suggest that the boundary between planktotrophy and lecithotrophy is frequently crossed and that this transition is reversible. Ecological changes represent the crossing of a functional threshold. Only after crossing the threshold, do larvae experience qualitatively different selective pressures that can lead to subsequent changes in morphology and development. Two different changes have occurred in the type of morphogenesis: the simplification of larval morphology that is associated with obligate (nonfeeding) lecithotrophy and the loss of the larval body plan in the evolution from indirect to direct development. It is the modification of morphology independent of the ecological changes that requires alterations in developmental processes, constrains evolutionary options, imposes irreversibility, and establishes the discrete nature of larval patterns in marine invertebrates.     

温度对马铃薯甲虫生长发育的影响

为了进一步明确温度对马铃薯甲虫Leptinotarsa decemlineata (Say)生长发育的影响, 在恒温条件, 研究了温度对马铃薯甲虫实验种群生长发育的影响。结果表明：温度对马铃薯甲虫各虫态的发育历期、存活率及种群的繁殖力有显著的影响。发育历期随温度的升高而缩短, 发育速率与温度呈显著的正相关。马铃薯甲虫世代存活率由大到小的顺序为27℃＞23℃＞19℃＞31℃＞15℃。27℃时成虫的产卵量最高, 单雌平均卵量为729.7粒;其次为23℃, 单雌平均卵量为639.2粒。并测得马铃薯甲虫卵、1龄幼虫、2龄幼虫、3龄幼虫、4龄幼虫、幼虫期、蛹期及全世代的发育起点温度分别为9.14, 10.50, 8.17, 10.28, 9.04, 9.59, 10.23和10.90℃, 有效积温分别为73.26, 43.22, 39.23, 34.05, 161.97, 273.02, 100.38和542.58日·度。据此认为温度对马铃薯甲虫实验种群的生长发育、存活和繁殖影响明显。     

温度对胡萝卜微管蚜生长发育繁殖的影响

【目标】明确温度对胡萝卜微管蚜Semiaphis heraclei(Takahashi)生长发育和繁殖的影响。【方法】在室内5个温度梯度下(19、22、25、28、31℃)观察并比较胡萝卜微管蚜的发育历期、存活率、存活寿命及产仔量。【结果】在19?31℃范围内,胡萝卜微管蚜各龄期及完整世代的发育历期均随着温度的升高而缩短,完成一个世代分别需要22.17、17.13、12.57、10.03和7.83 d。温度与发育速率呈极显著相关(P<0.01,r>0.8),温度越高发育速率越快。胡萝卜微管蚜4个若蚜期和世代的发育起点分别为14.15、13.87、13.64、15.06、12.92℃,有效积温分别为29.98、29.30、28.54、24.00和144.21日?度。建立了胡萝卜微管蚜各个发育阶段的历期预测式。在5个恒温下胡萝卜微管蚜1?4龄及世代的存活率随温度的变化而小幅波动。在19?31℃范围内,该蚜虫世代的存活率分别为40.0%、52.5%、62.5%、60.0%和47.5%。25℃该蚜虫的存活率相对较高,1?4龄及世代的存活率分别为82.5%、81.8%、92.6%、100%和62.5%。在相同温度条件下该蚜虫不同发育阶段的存活率存在差异。存活寿命随着温度的升高而缩短,19℃时寿命最长,为33.30 d,31℃时寿命最短,为15.40 d。产仔期随着温度的升高而缩短,19℃时产仔期为11.13 d,31℃时为7.57 d。在19?31℃范围内,该蚜虫的单雌产仔总量分别为26.33、27.93、32.53、27.13和17.93头。【结论】温度是影响胡萝卜微管蚜的生长发育、存活和繁殖的重要因素。25℃时胡萝卜微管蚜各龄期及世代的发育历期较短,存活率较高,单雌产仔总量最大。25℃较适合该蚜虫生长发育和繁殖。     

Species‐specificity of ethylene glycol‐induced developmental toxicity: toxicokinetic and whole embryo culture studies in the rabbit

High‐dose gavage exposure to ethylene glycol (EG) is teratogenic in rats, but not rabbits. To investigate the reason for this species difference, toxicokinetic and whole embryo culture (WEC) studies were conducted in gestation day 9 New Zealand White rabbits, and the data compared to very similar data previously generated in pregnant rats. In the toxicokinetic study, maximal levels of unchanged EG in rabbits were comparable to those reported for rats. However, maximal levels of EG's teratogenic metabolite, glycolic acid (GA), in rabbit maternal blood and embryo were only 46% and 10% of the respective levels in rats. The toxicokinetic profile suggested that the lower GA levels in rabbits were due to a slower rate of maternal metabolism of EG to GA, slow uptake of GA into the yolk sac cavity fluid which surrounds the embryo, and negligible transfer via the visceral yolk sac (VYS) placenta. In the WEC study, exposure of rabbit conceptuses to high concentrations (≤12.5 mM) of GA was without effect, which contrasts with reported effects in rat WEC at ≥3 mM. Overall, these data implicate toxicokinetics as an important factor underlying the species difference, although intrinsic insensitivity of the rabbit embryo might also be involved. Integration of these findings with published human data suggest that the rabbit is the more relevant model for human EG exposure, based on the negligible role of the rabbit VYS in placental transfer (humans lack a VYS) and similar rates of EG metabolism and extraembryonic fluid turnover. Birth Defects Res (Part B) 2008. © 2008 Wiley‐Liss, Inc.     

温度对沙蒿金叶甲生长发育和繁殖的影响

为了明确温度对沙蒿金叶甲Chrysolina aeruginosa Faldermann生长发育和繁殖的影响, 本实验在恒温条件（13, 18, 23, 28和33℃）下, 以白沙蒿Artemisia sphaerocephala Krasch为寄主植物, 研究了温度对沙蒿金叶甲实验种群生长发育和繁殖的影响。结果表明： 温度对沙蒿金叶甲各虫态的发育历期、 存活率以及种群繁殖力有显著影响。在13～28℃范围内, 各虫态的发育历期均随温度的升高而缩短, 发育速率与温度呈显著正相关。但是, 当温度上升至33℃时, 幼虫和蛹生长发育受到抑制, 其幼虫发育历期与18℃, 23℃和28℃下相比延长并达到了显著差异水平(P<0.05), 成虫不能羽化出土; 低温影响沙蒿金叶甲卵的存活率, 高温影响其蛹的存活率, 说明温度过高或过低均不利于沙蒿金叶甲的生长发育。成虫产卵量随环境温度变化的大小顺序为28℃＞23℃＞18℃＞13℃, 并存在极显著差异(P<0.01)。由直接最优法计算得到沙蒿金叶甲卵期、 1-3龄幼虫期、 4龄幼虫+蛹期的发育起点温度分别为9.72℃, 7.11℃和8.77℃, 有效积温依次为115.36, 441.91和448.40日·度。研究结果为沙蒿金叶甲发生期的预测预报提供了基础参考数据, 对指导生产实践有实际的应用意义。     

寄生植物锁阳茎的发育解剖学研究

锁阳茎的初生分生组织由原表皮、基本分生组织以及在基本分生组织中呈波浪式环状排列的原形成层束组成。茎的增粗是由于呈波浪式环状排列的维管束,其“波浪”上下幅度逐渐增大,即从“浪”的基部到“浪”顶端维管束数目由4个逐渐增加到10－12个。维管束数目不断增加是由于：（1）由髓射线薄壁细胞反分化产生分生组织束,分生组织束活动产生新的维管束;（2）维管束中分化出一列或几列薄壁细胞,导致该维管束被分化出的薄壁细胞分成2－3个独立的维管束。