Differential growth

差异增长
  • 文章类型: Journal Article
    目的:描述青少年特发性脊柱侧凸(AIS)患者脊柱生长的生理学。
    方法:关注生长机制的文献叙事回顾。
    结果:在他具有里程碑意义的出版物《成长与形式》中,阿西·汤普森写道,有机体的解剖结构反映了它所承受的力量。这意味着机械力是组织形状的基础,器官和有机体,无论是健康还是患病。AIS被称为特发性,因为变形的根本原因是未知的,尽管许多因素是相关的。最终,然而,任何畸形都是由于机械力。长期以来,研究表明,脊柱侧弯的典型弯曲和旋转可能是由于椎骨和椎间盘的生长速度快于附着在其上的韧带。这提出了一个问题,为什么在AIS中,韧带不能跟上脊柱生长的速度。AIS患者的脊柱以各种方式偏离健康的脊柱。增长较晚但更快,导致较高的椎骨和椎间盘。椎骨密度较低,这表明脊髓压迫较少。这也保留了脊索细胞和髓核中的肿胀压力。较少的脊髓压迫是由于有限的肌肉活动,低肌肉质量确实是AIS患者较低体重指数(BMI)的基础。因此,AIS棘生长得更快,因为脊柱压迫抵消了生长力(休特-沃尔克曼定律)。韧带由通过张力生长的胶原纤维组成,纤维状滑动和交联的重塑。动态负荷和雌激素等激素可增强生长和重塑。然而,他们反对静态加载。
    结论:脊柱伸长增加和韧带生长减少导致差异应变和脊柱侧弯变形的恶性循环。认识到有助于差异生长的物理和生物学线索可以更早地诊断AIS并预防处于危险中的儿童。
    OBJECTIVE: To describe the physiology of spinal growth in patients with adolescent idiopathic scoliosis (AIS).
    METHODS: Narrative review of the literature with a focus on mechanisms of growth.
    RESULTS: In his landmark publication On Growth and Form, D\'Arcy Thompson wrote that the anatomy of an organism reflects the forces it is subjected to. This means that mechanical forces underlie the shape of tissues, organs and organisms, whether healthy or diseased. AIS is called idiopathic because the underlying cause of the deformation is unknown, although many factors are  associated. Eventually, however, any deformity is due to mechanical forces. It has long been shown that the typical curvature and rotation of the scoliotic spine could result from vertebrae and intervertebral discs growing faster than the ligaments attached to them. This raises the question why in AIS the ligaments do not keep up with the speed of spinal growth. The spine of an AIS patient deviates from healthy spines in various ways. Growth is later but faster, resulting in higher vertebrae and intervertebral discs. Vertebral bone density is lower, which suggests  less spinal compression. This also preserves the notochordal cells and the swelling pressure in the nucleus pulposus. Less spinal compression is due to limited muscular activity, and low muscle mass indeed underlies the lower body mass index (BMI) in AIS patients. Thus, AIS spines grow faster because there is less spinal compression that counteracts the force of growth (Hueter-Volkmann Law). Ligaments consist of collagen fibres that grow by tension, fibrillar sliding and the remodelling of cross-links. Growth and remodelling are enhanced by dynamic loading and by hormones like estrogen. However, they are opposed by static loading.
    CONCLUSIONS: Increased spinal elongation and reduced ligamental growth result in differential strain and a vicious circle of scoliotic deformation. Recognising the physical and biological cues that contribute to differential growth  allows earlier diagnosis of AIS and prevention in children at risk.
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  • 文章类型: Journal Article
    本文关注的是增长驱动的形状规划问题,这涉及确定生长张量,该张量可以在达到给定目标形状的超弹性体上产生变形。我们考虑全球兼容增长的两种情况,其中生长张量是未变形域上的变形梯度,和不相容的,这抛弃了这样的假设。我们在超弹性最优控制理论的框架内提出了问题。Hausdorff距离用于量化形状之间的差异;致动的复杂性也被纳入成本函数中。边界条件和外部载荷在州法中是允许的,从而扩展了以前的工作,无压力假设被证明是必不可少的。然后进行严格的数学分析以证明问题的适当性。使用基于梯度的优化算法执行数值逼近。在这一部分中,我们的主要目标是展示将逆技术应用于该问题的数值逼近的可能性,这使我们能够解决比分析方法所涵盖的情况更通用的情况。梁状和壳型几何形状的一些数值实验说明了所提出的数值方案的性能。
    This paper is concerned with the growth-driven shape-programming problem, which involves determining a growth tensor that can produce a deformation on a hyperelastic body reaching a given target shape. We consider the two cases of globally compatible growth, where the growth tensor is a deformation gradient over the undeformed domain, and the incompatible one, which discards such hypothesis. We formulate the problem within the framework of optimal control theory in hyperelasticity. The Hausdorff distance is used to quantify dissimilarities between shapes; the complexity of the actuation is incorporated in the cost functional as well. Boundary conditions and external loads are allowed in the state law, thus extending previous works where the stress-free hypothesis turns out to be essential. A rigorous mathematical analysis is then carried out to prove the well-posedness of the problem. The numerical approximation is performed using gradient-based optimisation algorithms. Our main goal in this part is to show the possibility to apply inverse techniques for the numerical approximation of this problem, which allows us to address more generic situations than those covered by analytical approaches. Several numerical experiments for beam-like and shell-type geometries illustrate the performance of the proposed numerical scheme.
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  • 文章类型: Journal Article
    背景:前庭aqeductusventuli(渡槽)被认为与胚胎和成人的球囊相连。然而,在胚胎中,众所周知,囊和囊广泛交流,以提供共同的内淋巴空间“心房”。
    方法:使用来自五个胚胎的矢状组织学切片(冠-臀部长度或CRL,14-21mm),9个早期胎儿(CRL24-35mm)和12个中期和近期胎儿(CRL82-272mm),我们重温了人耳渡槽的发展和成长。
    结果:作为渡槽的前下延续,中庭呈现出厚厚的管状外观,但很快就分成了多个鸿沟。大多数的海湾对应于半圆形管道的壶腹,而前中下角的一个鸿沟对应于未来的囊。值得注意的是,在14个胚胎和早期胎儿中有8个,渡槽在前(上)或后半规管的原始壶腹附近的囊处终止。相反,CRL21mm的胚胎是最小的标本,其中渡槽与海湾状囊相连。在中期和近期,不断增长的外淋巴空间将渡槽与导管分开,并似乎将渡槽推向囊。胚胎位于上方的囊和位于下方的囊之间发生了地形变化,从而在成人中形成了前后排列。
    结论:因此,导水管的前庭端最有可能在6-8周时从囊向球囊向前迁移,这可能是由于内皮的差异生长。先前对胚胎渡槽的重建可能受到成人形态的影响。
    BACKGROUND: The aqeductus vestibuli (aqueduct) is believed to connect to the saccule in embryos and adults. However, in embryos, the saccule and utricle are known to communicate widely to provide a common endolymph space \"atrium\".
    METHODS: Using sagittal histological sections from five embryos (crown-rump length or CRL, 14-21 mm), nine early fetuses (CRL 24-35 mm) and 12 midterm and near-term fetuses (CRL 82-272 mm), we revisited the development and growth of the human ear aqueduct.
    RESULTS: The atrium took on a thick tube-like appearance as an antero-inferior continuation of the aqueduct, but soon divided into multiple gulfs. Most of the gulfs corresponded to the ampullae of semicircular ducts, while one gulf at the antero-medio-inferior corner corresponded to the future saccule. Notably, in eight of the 14 embryos and early fetuses, the aqueduct ended at the utricle near the primitive ampulla of the anterior (superior) or posterior semicircular duct. Conversely, an embryo of CRL 21 mm was the smallest specimen in which the aqueduct joined the gulf-like saccule. At midterm and near-term, the growing perilymph space separated the aqueduct from the utricle and appeared to push the aqueduct toward the saccule. A topographical change occurred between the embryonic superiorly located utricle and the inferiorly-located saccule to create the antero-posterior arrangement in adults.
    CONCLUSIONS: Consequently, the vestibular end of the aqueduct was most likely to migrate anteriorly from the utricle to the saccule at 6-8 weeks possibly due to differential growth of the endothelium. Previous reconstructions of the embryonic aqueduct might be biased by the adult morphology.
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  • 文章类型: Journal Article
    根尖钩是双子叶种子在黑暗中发芽时在下胚轴上部形成的简单弯曲结构。钩结构是短暂的,但对于幼苗在土壤出苗期间的生存至关重要,因为它有效地保护了脆弱的茎尖免受机械伤害。作为研究植物差异生长的极好模型系统,根尖钩早在达尔文时代就吸引了植物学家,并且在形态和分子水平上都取得了重大进展,以了解根尖钩发育是如何调节的。这里,我们将主要总结这两个方面的研究进展。我们还将简要比较种子萌发初期顶端钩和下胚轴重力弯曲之间的生长动力学,目的是就他们之间的联系达成某种共识。最后,我们将概述根尖钩发育的剩余问题和未来研究前景。本文受版权保护。保留所有权利。
    Apical hook is a simple curved structure formed at the upper part of hypocotyls when dicot seeds germinate in darkness. The hook structure is transient but essential for seedlings\' survival during soil emergence due to its efficient protection of the delicate shoot apex from mechanical injury. As a superb model system for studying plant differential growth, apical hook has fascinated botanists as early as the Darwin age, and significant advances have been achieved at both the morphological and molecular levels to understand how apical hook development is regulated. Here, we will mainly summarize the research progress at these two levels. We will also briefly compare the growth dynamics between apical hook and hypocotyl gravitropic bending at early seed germination phase, with the aim to deduce a certain consensus on their connections. Finally, we will outline the remaining questions and future research perspectives for apical hook development.
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  • 文章类型: Journal Article
    气管是一根长管,可以在喉和支气管之间通过空气。腹侧的C形软骨环使结构稳定。在它面向食道的背侧,可变形的平滑肌有助于食物在食道中通过。虽然沿背腹轴的对称断裂是很好理解的,导致转变为软骨环的周期性Sox9表达模式的分子机制仍然难以捉摸。这里,我们回顾了已经阐明的分子调控相互作用,并讨论可能的图案化机制。理解自组织的原则很重要,既可以定义生物医学干预措施,又可以实现组织工程。
    The trachea is a long tube that enables air passage between the larynx and the bronchi. C-shaped cartilage rings on the ventral side stabilise the structure. On its esophagus-facing dorsal side, deformable smooth muscle facilitates the passage of food in the esophagus. While the symmetry break along the dorsal-ventral axis is well understood, the molecular mechanism that results in the periodic Sox9 expression pattern that translates into the cartilage rings has remained elusive. Here, we review the molecular regulatory interactions that have been elucidated, and discuss possible patterning mechanisms. Understanding the principles of self-organisation is important, both to define biomedical interventions and to enable tissue engineering.
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  • 文章类型: Journal Article
    这项研究旨在确定(1)椎体束缚(VBT)是否在特发性脊柱侧凸患者的个体椎骨中产生不同的生长调节,(2)VBT是否改变圆盘形状,和(3)VBT是否影响脊柱总长度?
    包括接受主胸弯VBT治疗并至少2年随访的特发性脊柱侧凸患者。椎体和椎间盘被归类为无器械近端胸椎,仪器化的主胸,或无器械胸腰椎。在随后的X光片上测量每个椎骨和椎间盘的左侧和右侧高度,以评估差异生长。将T1-T12胸椎和T1-S1胸腰段生长速度与标准化参考数据进行比较。
    分析了51例患者(764个椎骨和807个椎间盘)。在2年的随访中,平均主曲线幅度从46°±11°提高到17°±11°。在MT椎骨中观察到差异生长,其中左侧/凹侧增加2.0±2.2mm,而右侧/凸侧增加1.5±2.3mm(p<0.001)。观察到所有光盘的高度变化差异,但在仪器化的MT光盘中最为明显,其中右侧/凸侧平均每个减少1.2毫米,与左侧/凹侧高度无明显变化相比。脊柱总生长速度与标准参考数据没有显着差异。
    椎体束缚限制了所设计的凸脊柱生长,同时允许凹形生长。曲线校正是由于椎体生长差异和凸起椎间盘高度降低所致。总体脊柱生长以预期的速率继续。
    IV级案例系列。
    This study aimed to determine (1) does vertebral body tethering (VBT) produce differential growth modulation in individual vertebrae in patients with idiopathic scoliosis, (2) does VBT change disc shape, and (3) does VBT affect total spine length?
    Patients with idiopathic scoliosis treated with VBT of the main thoracic curve and minimum 2-year follow-up were included. Vertebrae and discs were categorized as uninstrumented proximal thoracic, instrumented main thoracic, or uninstrumented thoracolumbar-lumbar. The left- and right-sided heights of each vertebra and disc were measured on subsequent radiographs to assess for differential growth. T1-T12 thoracic and T1-S1 thoracolumbar growth velocities were compared with standardized reference data.
    Fifty-one patients (764 vertebrae and 807 discs) were analyzed. The average major curve magnitude improved from 46° ± 11° to 17° ± 11° at 2-year follow-up. Differential growth was observed in MT vertebrae, in which the left/concave side grew 2.0 ± 2.2 mm compared with 1.5 ± 2.3 mm on the right/convex (tethered) side (p < 0.001). Differential height changes were observed for all discs, but were most pronounced in instrumented MT discs, in which the right/convex sides decreased by an average of 1.2 mm each, compared with no significant height change on the left/concave side. Total spinal growth velocities were not significantly different from standard reference data.
    Vertebral body tethering limits convex spinal growth as designed while permitting concave growth. Curve correction results from differential vertebral growth and decreased convex disc height. Overall spinal growth continues at the expected rate.
    Level IV case series.
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  • 文章类型: Journal Article
    The bending of epithelial tubes is a fundamental process in organ morphogenesis, driven by various multicellular behaviours. The cochlea in the mammalian inner ear is a representative example of spiral tissue architecture where the continuous bending of the duct is a fundamental component of its morphogenetic process. Although the cochlear duct morphogenesis has been studied by genetic approaches extensively, it is still unclear how the cochlear duct morphology is physically formed. Here, we report that nuclear behaviour changes are associated with the curvature of the pseudostratified epithelium during murine cochlear development. Two-photon live-cell imaging reveals that the nuclei shuttle between the luminal and basal edges of the cell is in phase with cell-cycle progression, known as interkinetic nuclear migration, in the flat region of the pseudostratified epithelium. However, the nuclei become stationary on the luminal side following mitosis in the curved region. Mathematical modelling together with perturbation experiments shows that this nuclear stalling facilitates luminal-basal differential growth within the epithelium, suggesting that the nuclear stalling would contribute to the bending of the pseudostratified epithelium during the cochlear duct development. The findings suggest a possible scenario of differential growth which sculpts the tissue shape, driven by collective nuclear dynamics.
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  • 文章类型: Journal Article
    植物根据环境条件将生物量分配给地上和地下器官。虽然广泛的模式是众所周知的,植物分配新生长的机制尚不清楚。生物量分配的建模方法广泛分为功能平衡型模型和基于机械的运输阻力型模型。我们在高光照和低光照下在分裂的根箱中种植早熟禾植物,高和低N供应,N相等或不相等地供应。我们的数据表明光照水平对根系质量的影响最强,N水平在控制芽质量方面更为重要。根系内生长的分配与韧皮部分区模型兼容。根质量分数受光和氮水平的影响,尽管在光照水平下,变化主要是由于芽生长的变化,根质量保持相对不变。在低光照条件下,植物表现出增加的比叶面积,大概是为了补偿低光水平。在后续实验中,我们表明,在弱光条件下,落叶可以抑制差异根的生长。我们的数据与运输阻力类型模型更兼容。
    Plants allocate biomass to above- and below-ground organs in response to environmental conditions. While the broad patterns are well-understood, the mechanisms by which plants allocate new growth remain unclear. Modeling approaches to biomass allocation broadly split into functional equilibrium type models and more mechanistically based transport resistance type models. We grew Poa annua plants in split root boxes under high and low light levels, high and low N supplies, with N supplied equally or unequally. Our data suggest that light level had the strongest effect on root mass, with N level being more important in controlling shoot mass. Allocation of growth within the root system was compatible with phloem partitioning models. The root mass fraction was affected by both light and N levels, although within light levels the changes were primarily due to changes in shoot growth, with root mass remaining relatively invariant. Under low light conditions, plants exhibited increased specific leaf area, presumably to compensate for low light levels. In a follow-up experiment, we showed that differential root growth could be suppressed by defoliation under low light conditions. Our data were more compatible with transport resistance type models.
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  • 文章类型: Journal Article
    3铰链回脊折叠是来自三个不同方向的回脊线的结合。以前的研究还没有探索这种三铰链回旋形成的可能机制,它们被保存在灵长类动物大脑中的物种中。我们开发了一个生物力学模型来模拟真实大脑上3铰链模式的形成,并确定在模型的某些区域如何形成特殊类型的3铰链模式。我们的计算和实验成像结果表明,生长和折叠后的大多数三次卷积和3铰链模式的确切位置是不可预测的,但它们有助于解释某些三铰链模式的位置和模式的一致性。白质中生长的纤维被认为是影响这些三铰链图案的位置和形状的决定因素。即使生长的纤维没有施加足够强的力来直接引导旋化,它们仍然可能播种异质生长曲线,导致在特定位置形成3铰链模式。两个正在生长的模型大脑之间的初始形态的微小差异可以导致折叠后3铰链模式的不同数量和位置。
    The 3-hinge gyral folding is the conjunction of gyrus crest lines from three different orientations. Previous studies have not explored the possible mechanisms of formation of such 3-hinge gyri, which are preserved across species in primate brains. We develop a biomechanical model to mimic the formation of 3-hinge patterns on a real brain and determine how special types of 3-hinge patterns form in certain areas of the model. Our computational and experimental imaging results show that most tertiary convolutions and exact locations of 3-hinge patterns after growth and folding are unpredictable, but they help explain the consistency of locations and patterns of certain 3-hinge patterns. Growing fibers within the white matter is posited as a determining factor to affect the location and shape of these 3-hinge patterns. Even if the growing fibers do not exert strong enough forces to guide gyrification directly, they still may seed a heterogeneous growth profile that leads to the formation of 3-hinge patterns in specific locations. A minor difference in initial morphology between two growing model brains can lead to distinct numbers and locations of 3-hinge patterns after folding.
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  • 文章类型: Journal Article
    我们提出了一个三维形态弹性杆模型,该模型能够描述由差异生长驱动的生长植物芽的形态发生。我们讨论了内生振荡器的演化规律,矫直机制,并重新定向到方向提示,例如受陨石雪崩动力学控制的重力反应。我们使用此模型来研究重力载荷引起的弹性偏转在环绕植物芽中的作用。我们证明,在没有内源性线索的情况下,摆动和圆形振荡随着达到临界长度而出现,从而表明由外源因素引发的不稳定性的发生。当还存在由于内生线索引起的振荡时,随着射击长度和其他生物力学特性的变化,它们的重量相对于与不稳定性相关的重量随时间变化。由于这两种振荡机制的同时发生,我们能够重现各种复杂的行为,包括摆线样图案,随着射击长度的增加,它们演变成圆形轨道,外生振荡的幅度成为主导。
    We present a three-dimensional morphoelastic rod model capable to describe the morphogenesis of growing plant shoots driven by differential growth. We discuss the evolution laws for endogenous oscillators, straightening mechanisms, and reorientations to directional cues, such as gravitropic reactions governed by the avalanche dynamics of statoliths. We use this model to investigate the role of elastic deflections due to gravity loading in circumnutating plant shoots. We show that, in the absence of endogenous cues, pendular and circular oscillations arise as a critical length is attained, thus suggesting the occurrence of an instability triggered by exogenous factors. When also oscillations due to endogenous cues are present, their weight relative to those associated with the instability varies in time as the shoot length and other biomechanical properties change. Thanks to the simultaneous occurrence of these two oscillatory mechanisms, we are able to reproduce a variety of complex behaviors, including trochoid-like patterns, which evolve into circular orbits as the shoot length increases, and the amplitude of the exogenous oscillations becomes dominant.
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