Organ Shape Research Articles

Epidermal restriction confers robustness to organ shapes.

The shape of comparable tissues and organs is consistent among individuals of a given species, but how this consistency or robustness is achieved remains an open question. The interaction between morphogenetic factors determines organ formation and subsequent shaping, which is ultimately a mechanical process. Using a computational approach, we show that the epidermal layer is essential for the robustness of organ geometry control. Specifically, proper epidermal restriction allows organ asymmetry maintenance, and the tensile epidermal layer is sufficient to suppress local variability in growth, leading to shape robustness. The model explains the enhanced organ shape variations in epidermal mutant plants. In addition, differences in the patterns of epidermal restriction may underlie the initial establishment of organ asymmetry. Our results show that epidermal restriction can answer the longstanding question of how cellular growth noise is averaged to produce precise organ shapes, and the findings also shed light on organ asymmetry establishment.

The diversity of the genus Aricidea (Polychaeta: Paraonidae) from the Sea of Marmara, with descriptions of two new species and two new records for the Mediterranean fauna.

This paper reports 13 Aricidea species from the Sea of Marmara. The Aricidea specimens were collected at 98 stations in the region in 2012 and 2013 from soft and hard substrata at depths ranging from 0 to 1200 m. Among the material, two new species, namely Aricidea (Acmira) katzmanni n. sp. and Aricidea (Acmira) pseudoassimilis n. sp., and 11 known species were discovered. Aricidea mirunekoa and A. bulbosa are new records for the Mediterranean fauna; A. (Acmira) annae and A. (Aricidea) wassi are new records for the Sea of Marmara. Aricidea (Acmira) katzmanni n. sp. is characterized by having a long, digitiform antenna; no notopodial papillae on the posterior part of the branchial region; and hook shaped modified neurochaetae with a strong hood and fragile arista. Aricidea (Acmira) pseudoassimilis n. sp. is characterized by having a short, digitiform antenna; interramal lobes between notopodia and neuropodia; hook-shaped modified neurochaeta that gets subterminally thinner with a rounded tip. As A. (Strelzovia) bulbosa has been only previously reported from the Gulf of Suez (Red Sea), it could be a new alien species for the Sea of Marmara and Mediterranean Sea. Based on the characters that have been overlooked so far, the subspecies A. suecica meridionalis and A. capensis bansei were raised to species level, A. meridionalis n. stat. and A. bansei n. stat. The present paper reports the usage of ciliary bands and slits on the prostomium and body, swellings/ridges in the branchial region, and the shape and distribution of sense organs in the taxonomy of Aricidea. All species were described and figured.

Epithelial organ shape is generated by patterned actomyosin contractility and maintained by the extracellular matrix.

Epithelial sheets define organ architecture during development. Here, we employed an iterative multiscale computational modeling and quantitative experimental approach to decouple direct and indirect effects of actomyosin-generated forces, nuclear positioning, extracellular matrix, and cell-cell adhesion in shaping Drosophila wing imaginal discs. Basally generated actomyosin forces generate epithelial bending of the wing disc pouch. Surprisingly, acute pharmacological inhibition of ROCK-driven actomyosin contractility does not impact the maintenance of tissue height or curved shape. Computational simulations show that ECM tautness provides only a minor contribution to modulating tissue shape. Instead, passive ECM pre-strain serves to maintain the shape independent from actomyosin contractility. These results provide general insight into how the subcellular forces are generated and maintained within individual cells to induce tissue curvature. Thus, the results suggest an important design principle of separable contributions from ECM prestrain and actomyosin tension during epithelial organogenesis and homeostasis.

HEARTBREAK Controls Post-translational Modification of INDEHISCENT to Regulate Fruit Morphology in Capsella

SummaryMorphological variation is the basis of natural diversity and adaptation. For example, angiosperms (flowering plants) evolved during the Cretaceous period more than 100 mya and quickly colonized terrestrial habitats [1]. A major reason for their astonishing success was the formation of fruits, which exist in a myriad of different shapes and sizes [2]. Evolution of organ shape is fueled by variation in expression patterns of regulatory genes causing changes in anisotropic cell expansion and division patterns [3, 4, 5]. However, the molecular mechanisms that alter the polarity of growth to generate novel shapes are largely unknown. The heart-shaped fruits produced by members of the Capsella genus comprise an anatomical novelty, making it particularly well suited for studies on morphological diversification [6, 7, 8]. Here, we show that post-translational modification of regulatory proteins provides a critical step in organ-shape formation. Our data reveal that the SUMO protease, HEARTBREAK (HTB), from Capsella rubella controls the activity of the key regulator of fruit development, INDEHISCENT (CrIND in C. rubella), via de-SUMOylation. This post-translational modification initiates a transduction pathway required to ensure precisely localized auxin biosynthesis, thereby facilitating anisotropic cell expansion to ultimately form the heart-shaped Capsella fruit. Therefore, although variation in the expression of key regulatory genes is known to be a primary driver in morphological evolution, our work demonstrates how other processes—such as post-translational modification of one such regulator—affects organ morphology.

Evolution and development of three highly specialized floral structures of bee-pollinated Phalaenopsis species

BackgroundVariation in shape and size of many floral organs is related to pollinators. Evolution of such organs is driven by duplication and modification of MADS-box and MYB transcription factors. We applied a combination of micro-morphological (SEM and micro 3D-CT scanning) and molecular techniques (transcriptome and RT-PCR analysis) to understand the evolution and development of the callus, stelidia and mentum, three highly specialized floral structures of orchids involved in pollination. Early stage and mature tissues were collected from flowers of the bee-pollinated Phalaenopsis equestris and Phalaenopsis pulcherrima, two species that differ in floral morphology: P. equestris has a large callus but short stelidia and no mentum, whereas P. pulcherrima has a small callus, but long stelidia and a pronounced mentum.ResultsOur results show the stelidia develop from early primordial stages, whereas the callus and mentum develop later. In combination, the micro 3D-CT scan analysis and gene expression analyses show that the callus is of mixed petaloid-staminodial origin, the stelidia of staminodial origin, and the mentum of mixed sepaloid-petaloid-staminodial origin. SEP clade 1 copies are expressed in the larger callus of P. equestris, whereas AP3 clade 1 and AGL6 clade 1 copies are expressed in the pronounced mentum and long stelidia of P. pulcherrima. AP3 clade 4, PI-, AGL6 clade 2 and PCF clade 1 copies might have a balancing role in callus and gynostemium development. There appears to be a trade-off between DIV clade 2 expression with SEP clade 1 expression in the callus, on the one hand, and with AP3 clade 1 and AGL6 clade 1 expression in the stelidia and mentum on the other.ConclusionsWe detected differential growth and expression of MADS box AP3/PI-like, AGL6-like and SEP-like, and MYB DIV-like gene copies in the callus, stelidia and mentum of two species of Phalaenopsis, of which these floral structures are very differently shaped and sized. Our study provides a first glimpse of the evolutionary developmental mechanisms driving adaptation of Phalaenopsis flowers to different pollinators by providing combined micro-morphological and molecular evidence for a possible sepaloid–petaloid–staminodial origin of the orchid mentum.

Gibberellin Enhances the Anisotropy of Cell Expansion in the Growth Zone of the Maize Leaf.

Although plant organ shapes are defined by spatio-temporal variations of directional tissue expansion, this is a little characterized aspect of organ growth regulation. Although it is well known that the plant hormone gibberellin increases the leaf length/with ratio, its effects on cell expansion in the growing leaf are largely unknown. To understand how variations in rate and anisotropy of growth establish the typical monocotelydonous leaf shape, we studied the leaf growth zone of maize (Zea mays) with a kinematic analysis of cell expansion in the three directions of growth: proximo-distal, medio-lateral, and dorso-ventral. To determine the effect of gibberellin, we compared a gibberellin-deficient dwarf3 mutant and the overproducing UBI::GA20OX-1 line with their wild types. We found that, as expected, longitudinal growth was dominant throughout the growth zone. The highest degree of anisotropy occurred in the division zone, where relative growth rates in width and thickness were almost zero. Growth anisotropy was smaller in the elongation zone, due to higher lateral and dorso-ventral growth rates. Growth in all directions stopped at the same position. Gibberellin increased the size of the growth zone and the degree of growth anisotropy by stimulating longitudinal growth rates. Inversely, the duration of expansion was negatively affected, so that mature cell length was unaffected, while width and height of cells were reduced. Our study provides a detailed insight in the dynamics of growth anisotropy in the maize leaf and demonstrates that gibberellin specifically stimulates longitudinal growth rates throughout the growth zone.

Multiple loci linked to inversions are associated with eye size variation in species of the Drosophila virilis phylad

The size and shape of organs is tightly controlled to achieve optimal function. Natural morphological variations often represent functional adaptations to an ever-changing environment. For instance, variation in head morphology is pervasive in insects and the underlying molecular basis is starting to be revealed in the Drosophila genus for species of the melanogaster group. However, it remains unclear whether similar diversifications are governed by similar or different molecular mechanisms over longer timescales. To address this issue, we used species of the virilis phylad because they have been diverging from D. melanogaster for at least 40 million years. Our comprehensive morphological survey revealed remarkable differences in eye size and head shape among these species with D. novamexicana having the smallest eyes and southern D. americana populations having the largest eyes. We show that the genetic architecture underlying eye size variation is complex with multiple associated genetic variants located on most chromosomes. Our genome wide association study (GWAS) strongly suggests that some of the putative causative variants are associated with the presence of inversions. Indeed, northern populations of D. americana share derived inversions with D. novamexicana and they show smaller eyes compared to southern ones. Intriguingly, we observed a significant enrichment of genes involved in eye development on the 4th chromosome after intersecting chromosomal regions associated with phenotypic differences with those showing high differentiation among D. americana populations. We propose that variants associated with chromosomal inversions contribute to both intra- and interspecific variation in eye size among species of the virilis phylad.

Statistical shape analysis of tap roots: a methodological case study on laser scanned sugar beets

BackgroundThe efficient and robust statistical analysis of the shape of plant organs of different cultivars is an important investigation issue in plant breeding and enables a robust cultivar description within the breeding progress. Laserscanning is a highly accurate and high resolution technique to acquire the 3D shape of plant surfaces. The computation of a shape based principal component analysis (PCA) built on concepts from continuum mechanics has proven to be an effective tool for a qualitative and quantitative shape examination.ResultsThe shape based PCA was used for a statistical analysis of 140 sugar beet roots of different cultivars. The calculation of the mean sugar beet root shape and the description of the main variations was possible. Furthermore, unknown and individual tap roots could be attributed to their cultivar by means of a robust classification tool based on the PCA results.ConclusionThe method demonstrates that it is possible to identify principal modes of root shape variations automatically and to quantify associated variances out of laserscanned 3D sugar beet tap root models. The introduced approach is not limited to the 3D shape description by laser scanning. A transfer to 3D MRI or radar data is also conceivable.

3D printed spermathecae as experimental models to understand sperm dynamics in leaf beetles

BackgroundPostcopulatory mate choice occurs ubiquitously in the animal kingdom. However, it is usually a major challenge to visualise the process taking place in a body. This fact makes it difficult to understand the mechanisms of the process. By focusing on the shape of female sperm storage organs (spermathecae), we aimed to elucidate their functional morphology using six representative beetle species and to simulate sperm dynamics in artificial spermathecae with different structural features.ResultsMorphology and material gradients were studied using micro-computed tomography (μCT) and confocal laser scanning microscopy. This study shows a diversity of external and internal structures of the spermathecae among species. Despite the diversity, all species possess a common pumping region, which is composed of a sclerotised chamber, muscles and a resilin-enriched region. By focusing on the species Agelastica alni, whose spermatheca is relatively simple in shape with an internal protuberance, we simulated sperm dynamics by establishing a fabrication method to create enlarged, transparent, flexible and low-cost 3D models of biological structures based on μCT data. This experiment shows that the internal protuberance in the species functions as an efficient mixing device of stored sperm.ConclusionsThe observed spermathecal musculature implies that the sclerotised chamber of the spermatheca with muscles works as a pumping organ. Our fluid dynamics tests based on 3D printed spermathecae show that a tiny structural difference causes entirely different fluid dynamics in the spermatheca models. This result suggests that structural variations of the spermatheca strongly affect sperm dynamics. However, fluid dynamics tests still require essential measurements including sperm viscosity and the velocity of pumping cycles of the spermatheca.

An Improved Mask R-CNN Model for Multiorgan Segmentation

Medical image segmentation is a key topic in image processing and computer vision. Existing literature mainly focuses on single-organ segmentation. However, since maximizing the concentration of radiotherapy drugs in the target area with protecting the surrounding organs is essential for making effective radiotherapy plan, multiorgan segmentation has won more and more attention. An improved Mask R-CNN (region-based convolutional neural network) model is proposed for multiorgan segmentation to aid esophageal radiation treatment. Due to the fact that organ boundaries may be fuzzy and organ shapes are various, original Mask R-CNN works well on natural image segmentation while leaves something to be desired on the multiorgan segmentation task. Addressing it, the advantages of this method are threefold: (1) a ROI (region of interest) generation method is presented in the RPN (region proposal network) which is able to utilize multiscale semantic features. (2) A prebackground classification subnetwork is integrated to the original mask generation branch to improve the precision of multiorgan segmentation. (3) 4341 CT images of 44 patients are collected and annotated to evaluate the proposed method. Additionally, extensive experiments on the collected dataset demonstrate that the proposed method can segment the heart, right lung, left lung, planning target volume (PTV), and clinical target volume (CTV) accurately and efficiently. Specifically, less than 5% of the cases were missed detection or false detection on the test set, which shows a great potential for real clinical usage.

Live Tissue Imaging Sheds Light on Cell Level Events During Ectodermal Organ Development.

Embryonic development of ectodermal organs involves a very dynamic range of cellular events and, therefore, requires advanced techniques to visualize them. Ectodermal organogenesis proceeds in well-defined sequential stages mediated by tissue interactions. Different ectodermal organs feature shared morphological characteristics, which are regulated by conserved and reiterative signaling pathways. A wealth of genetic information on the expression patterns and interactions of specific signaling pathways has accumulated over the years. However, the conventional developmental biology methods have mainly relied on two-dimensional tissue histological analyses at fixed time points limiting the possibilities to follow the processes in real time on a single cell resolution. This has complicated the interpretation of cause and effect relationships and mechanisms of the successive events. Whole-mount tissue live imaging approaches are now revealing how reshaping of the epithelial sheet for the initial placodal thickening, budding morphogenesis and beyond, involve coordinated four dimensional changes in cell shapes, well-orchestrated cell movements and specific cell proliferation and apoptosis patterns. It is becoming evident that the interpretation of the reiterative morphogenic signals takes place dynamically at the cellular level. Depending on the context, location, and timing they drive different cell fate choices and cellular interactions regulating a pattern of behaviors that ultimately defines organ shapes and sizes. Here we review how new tissue models, advances in 3D and live tissue imaging techniques have brought new understanding on the cell level behaviors that contribute to the highly dynamic stages of morphogenesis in teeth, hair and related ectodermal organs during development, and in dysplasia contexts.

Holistic multitask regression network for multiapplication shape regression segmentation

A holistic multitask regression approach was implemented to tackle the limitations of clinical image analysis. Standard practice requires identifying multiple anatomic structures in multiple planes from multiple anatomic regions using multiple modalities. The proposed novel holistic multitask regression network (HMR-Net) formulates organ segmentation as a multitask learning problem. Multitask learning leverages the strength of joint task problem solving from capturing task correlations. HMR-Net performs multitask regression by estimating an organ's class, regional location, and precise contour coordinates. The estimation of each coordinate point also corresponds to another regression task. HMR-Net leverages hierarchical multiscale and fused organ features to handle nonlinear relationships between image appearance and distinct organ properties. Simultaneously, holistic shape information is captured by encoding coordinate correlations. The multitask pipeline enables the capturing of holistic organ information (e.g. class, location, shape) to perform shape regression for medical image segmentation. HMR-Net was validated on eight representative datasets obtained from a total of 222 subjects. A mean average precision and dice score reaching up to 0.81 and 0.93, respectively, was achieved on the representative multiapplication database. The generalized model demonstrates comparable or superior performance compared to state-of-the-art algorithms. The high-performance accuracy demonstrates our model as an effective general framework to perform organ shape regression in multiple applications. This method was proven to provide high-contrast sensitivity to delineate even the smallest and oddly shaped organs. HMR-Net's flexible framework holds great potential in providing a fully automatic preliminary analysis for multiple types of medical images.

Model-based registration for pneumothorax deformation analysis using intraoperative cone-beam CT images.

Because the lung deforms during surgery because of pneumothorax, it is important to be able to track the location of a tumor. Deformation of the whole lung can be estimated using intraoperative cone-beam CT (CBCT) images. In this study, we used deformable mesh registration methods for paired CBCT images in the inflated and deflated states, and analyzed their deformation. We proposed a deformable mesh registration framework for deformations of partial organ shapes involving large deformation and rotation. Experimental results showed that the proposed methods reduced errors in point-to-point correspondence. As a result of registration using surgical clips placed on the lung surface during imaging, it was confirmed that an average error of 3.9 mm occurred in eight cases. The result of analysis showed that both tissue rotation and contraction had large effects on displacement.

Stable and efficient transformation of apple

Apple is one of precious fruit crop grown in temperate zone. In the post genomic era, the analysis of gene functions in horticultural crops such as apple is required for agricultural utilization. For analysis of such crops, the protocol establishment of tissue culture and transformation is essential. Although transformation efficiency in family Rosaceae is generally very low, some cultivars of Malus species have high transformation ability. Apple cultivars are usually clonally propagated by grafting on rootstocks, which can affect fruit quality and maturity and scion productivity. Apple rootstock cultivar Japan Morioka 2 (JM2) was produced at the Division of Apple Research, Institute of Fruit and Tea Science, NARO, in Japan. JM2, which was developed for dwarfing scions and improving disease resistance, is easily propagated by hardwood cutting. Furthermore, JM2 can be stably transformed at a high efficiency, which is better than other JM series rootstocks derived from the same parent. Leaflets of cultured shoots of JM2 have been transformed using Agrobacterium (Rhizobium) with a transducing gene. In this article, the JM2 transformation protocol is introduced in detail. Various genes and promoters have been confirmed to function as expected, with the resultant transformants exhibiting specific staining and fluorescent signals, and modified floral organ shapes, precious blooming and other characteristics. JM2 is thus a useful rootstock material for the enhancement of genetic research on apple and its relatives.

Spatiotemporal Pattern of Ectopic Cell Divisions Contribute to Mis-Shaped Phenotype of Primary and Lateral Roots of katanin1 Mutant.

Pattern formation, cell proliferation, and directional cell growth, are driving factors of plant organ shape, size, and overall vegetative development. The establishment of vegetative morphogenesis strongly depends on spatiotemporal control and synchronization of formative and proliferative cell division patterns. In this context, the progression of cell division and the regulation of cell division plane orientation are defined by molecular mechanisms converging to the proper positioning and temporal reorganization of microtubule arrays such as the preprophase microtubule band, the mitotic spindle and the cytokinetic phragmoplast. By focusing on the tractable example of primary root development and lateral root emergence in Arabidopsis thaliana, genetic studies have highlighted the importance of mechanisms underlying microtubule reorganization in the establishment of the root system. In this regard, severe alterations of root growth, and development found in extensively studied katanin1 mutants of A. thaliana (fra2, lue1, and ktn1-2), were previously attributed to defective rearrangements of cortical microtubules and aberrant cell division plane reorientation. How KATANIN1-mediated microtubule severing contributes to tissue patterning and organ morphogenesis, ultimately leading to anisotropy in microtubule organization is a trending topic under vigorous investigation. Here we addressed this issue during root development, using advanced light-sheet fluorescence microscopy (LSFM) and long-term imaging of ktn1-2 mutant expressing the GFP-TUA6 microtubule marker. This method allowed spatial and temporal monitoring of cell division patterns in growing roots. Analysis of acquired multidimensional data sets revealed the occurrence of ectopic cell divisions in various tissues including the calyptrogen and the protoxylem of the main root, as well as in lateral root primordia. Notably the ktn1-2 mutant exhibited excessive longitudinal cell divisions (parallel to the root axis) at ectopic positions. This suggested that changes in the cell division pattern and the occurrence of ectopic cell divisions contributed significantly to pleiotropic root phenotypes of ktn1-2 mutant. LSFM provided evidence that KATANIN1 is required for the spatiotemporal control of cell divisions and establishment of tissue patterns in living A. thaliana roots.

Evolution of the hectocotylus in Sepiolinae (Cephalopoda: Sepiolidae) and description of four new genera

The subfamily Sepiolinae (Mollusca: Cephalopoda: Sepiolidae), currently containing the genera Sepiola Leach, 1817, Euprymna Steenstrup, 1887, Inioteuthis Verrill, 1881, Rondeletiola Naef, 1921 and Sepietta Naef, 1912, is characterized by the hectocotylization of the left dorsal arm, i.e., its transformation into a copulatory organ thanks to modifications of sucker/pedicel elements. The hectocotylus morphology varies to a great extent across genera and species. In particular, one to several pedicels in its proximal third lose their sucker and become highly and diversely modified in shape to constitute a copulatory apparatus. An evolutionary gradient was observed in the copulatory apparatus morphology, from the simple modification into a papilla of just one pedicel from the third element of the ventral sucker row (some nominal species of Euprymna) to a quite complex structure involving several variously modified pedicels from both the ventral and dorsal sucker rows (Inioteuthis). In some species, elements in the distal portion of the hectocotylus may also be highly modified, such as the columnar suckers in Euprymna. The hectocotylian diversity allows to distinguish nine groups of species that do not match the current generic subdivision of Sepiolinae. Additional morphological characters (number of sucker rows on arms, female bursa copulatrix, occurrence and shape of visceral light organs, etc.) corroborate the subdivision of Sepiolinae into nine subtaxa, i.e., genera. Accordingly, a cladogram is drawn to describe the possible phylogenetic relationships among the nine clades. To comply with these results, all current genera are redefined and four new genera are described, namely Adinaefiola gen. nov., Boletzkyola gen. nov., Eumandya gen. nov. and Lusepiola gen. nov.

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Robust organ size requires robust timing of initiation orchestrated by focused auxin and cytokinin signalling.

Organ size and shape are precisely regulated to ensure proper function. The four sepals in each Arabidopsis thaliana flower must maintain the same size throughout their growth to enclose and protect the developing bud. Here we show that DEVELOPMENT RELATED MYB-LIKE1 (DRMY1) is required for both timing of organ initiation and proper growth, leading to robust sepal size in Arabidopsis. Within each drmy1 flower, the initiation of some sepals is variably delayed. Late-initiating sepals in drmy1 mutants remain smaller throughout development resulting in variability in sepal size. DRMY1 focuses the spatiotemporal signaling patterns of the plant hormones auxin and cytokinin, which jointly control the timing of sepal initiation. Our findings demonstrate that timing of organ initiation together with growth and maturation contribute to robust organ size.

Tailor-made 3D dosimeter using 3D printing technology

This article reports initial results of fabricating a tailor-made 3D dosimeter using radiochromic material and a 3D printer. The 3D dose distribution is easily visualized and quantitated as the transparent polymers become colored after irradiation. With this technology, we propose a novel method that uses 3D dosimetry to precisely copy the shape of human organs with our 3D printed dosimeter and help provide more accurate and safe radiotherapy.

Regulation of size and scale in vertebrate spinal cord development.

All vertebrates have a spinal cord with dimensions and shape specific to their species. Yet how species‐specific organ size and shape are achieved is a fundamental unresolved question in biology. The formation and sculpting of organs begins during embryonic development. As it develops, the spinal cord extends in anterior–posterior direction in synchrony with the overall growth of the body. The dorsoventral (DV) and apicobasal lengths of the spinal cord neuroepithelium also change, while at the same time a characteristic pattern of neural progenitor subtypes along the DV axis is established and elaborated. At the basis of these changes in tissue size and shape are biophysical determinants, such as the change in cell number, cell size and shape, and anisotropic tissue growth. These processes are controlled by global tissue‐scale regulators, such as morphogen signaling gradients as well as mechanical forces. Current challenges in the field are to uncover how these tissue‐scale regulatory mechanisms are translated to the cellular and molecular level, and how regulation of distinct cellular processes gives rise to an overall defined size. Addressing these questions will help not only to achieve a better understanding of how size is controlled, but also of how tissue size is coordinated with the specification of pattern.This article is categorized under:Establishment of Spatial and Temporal Patterns > Regulation of Size, Proportion, and TimingSignaling Pathways > Global Signaling MechanismsNervous System Development > Vertebrates: General Principles