Division Patterns Research Articles

Co‐contributorship network and division of labor in individual scientific collaborations

AbstractCollaborations are pervasive in current science. Collaborations have been studied and encouraged in many disciplines. However, little is known about how a team really functions from the detailed division of labor within. In this research, we investigate the patterns of scientific collaboration and division of labor within individual scholarly articles by analyzing their co‐contributorship networks. Co‐contributorship networks are constructed by performing the one‐mode projection of the author–task bipartite networks obtained from 138,787 articles published in PLoS journals. Given an article, we define 3 types of contributors: Specialists, Team‐players, and Versatiles. Specialists are those who contribute to all their tasks alone; team‐players are those who contribute to every task with other collaborators; and versatiles are those who do both. We find that team‐players are the majority and they tend to contribute to the 5 most common tasks as expected, such as “data analysis” and “performing experiments.” The specialists and versatiles are more prevalent than expected by our designed 2 null models. Versatiles tend to be senior authors associated with funding and supervision. Specialists are associated with 2 contrasting roles: the supervising role as team leaders or marginal and specialized contributors.

Adult neurogenesis and gliogenesis in the dorsal and ventral canine hippocampus.

Dentate gyrus (DG) of the mammalian hippocampus gives rise to new neurons and astrocytes all through adulthood. Canine hippocampus presents many similarities in fetal development, anatomy, and physiology with human hippocampus, establishing canines as excellent animal models for the study of adult neurogenesis. In the present study, BrdU-dated cells of the structurally and functionally dissociated dorsal (dDG) and ventral (vDG) adult canine DG were comparatively examined over a period of 30 days. Each part's neurogenic potential, radial glia-like neural stem cells (NSCs) proliferation and differentiation, migration, and maturation of their progenies were evaluated at 2, 5, 14, and 30 days post BrdU administration, with the use of selected markers (glial fibrillary acidic protein, doublecortin, calretinin and calbindin). Co-staining of BrdU+ cells with NeuN or S100B permitted the parallel study of the ongoing neurogenesis and gliogenesis. Our findings reveal the comparatively higher populations of residing granule cells, proliferating NSCs and BrdU+ neurons in the dDG, whereas newborn neurons of the vDG showed a prolonged differentiation, migration, and maturation. Newborn astrocytes were found all along the dorso-ventral axis, counting however for only 11% of newborn cell population. Comparative evaluation of adult canine and rat neurogenesis revealed significant differences in the distribution of resident and newborn granule cells along the dorso-ventral axis, division pattern of adult NSCs, maturation time plan of newborn neurons, and ongoing gliogenesis. Concluding, spatial and temporal features of adult canine neurogenesis are similar to that of other gyrencephalic species, including humans, and justify the comparative examination of adult neurogenesis across mammalian species.

Biology of the Caenorhabditis elegans Germline Stem Cell System.

Stem cell systems regulate tissue development and maintenance. The germline stem cell system is essential for animal reproduction, controlling both the timing and number of progeny through its influence on gamete production. In this review, we first draw general comparisons to stem cell systems in other organisms, and then present our current understanding of the germline stem cell system in Caenorhabditis elegans In contrast to stereotypic somatic development and cell number stasis of adult somatic cells in C. elegans, the germline stem cell system has a variable division pattern, and the system differs between larval development, early adult peak reproduction and age-related decline. We discuss the cell and developmental biology of the stem cell system and the Notch regulated genetic network that controls the key decision between the stem cell fate and meiotic development, as it occurs under optimal laboratory conditions in adult and larval stages. We then discuss alterations of the stem cell system in response to environmental perturbations and aging. A recurring distinction is between processes that control stem cell fate and those that control cell cycle regulation. C. elegans is a powerful model for understanding germline stem cells and stem cell biology.

Caulobactercrescentus β sliding clamp employs a noncanonical regulatory model of DNA replication.

The eubacterial β sliding clamp (DnaN) plays a crucial role in DNA metabolism through direct interactions with DNA, polymerases, and a variety of protein factors. A canonical protein-DnaN interaction has been identified in Escherichiacoli and some other species, during which protein partners are tethered into the conserved canonical hydrophobic crevice of DnaN via the consensus β-binding motif. Caulobactercrescentus is an excellent research model for use in the investigation of DNA replication and cell-cycle regulation due to its unique asymmetric cell division pattern with restricted replication initiation; however, little is known about the specific features of C.crescentus DnaN (CcDnaN). Here, we report a significant divergence in the association of CcDnaN with proteins based on docking analysis and crystal structures that show that the β-binding motifs of its protein partners bind a novel pocket instead of the canonical site. Pull-down and isothermal titration calorimetry results revealed that mutations within the novel pocket disrupt protein-CcDnaN interactions. It was also shown by replication and regulatory inactivation of DnaA assays that mediation of protein interaction by the novel pocket is closely related to the performance of CcDnaN during replication and the DnaN-mediated regulation process. Moreover, assessments of clamp competition showed that DNA does not compete with protein partners when binding to the novel pocket. Overall, our structural and biochemical analyses provide strong evidence that CcDnaN employs a noncanonical protein association pattern.

C. elegans Runx/CBFβ suppresses POP-1 TCF to convert asymmetric to proliferative division of stem cell-like seam cells.

ABSTRACTA correct balance between proliferative and asymmetric cell divisions underlies normal development, stem cell maintenance and tissue homeostasis. What determines whether cells undergo symmetric or asymmetric cell division is poorly understood. To gain insight into the mechanisms involved, we studied the stem cell-like seam cells in the Caenorhabditis elegans epidermis. Seam cells go through a reproducible pattern of asymmetric divisions, instructed by divergent canonical Wnt/β-catenin signaling, and symmetric divisions that increase the seam cell number. Using time-lapse fluorescence microscopy we observed that symmetric cell divisions maintain asymmetric localization of Wnt/β-catenin pathway components. Our observations, based on lineage-specific knockout and GFP-tagging of endogenous pop-1, support the model that POP-1TCF induces differentiation at a high nuclear level, whereas low nuclear POP-1 promotes seam cell self-renewal. Before symmetric division, the transcriptional regulator RNT-1Runx and cofactor BRO-1CBFβ temporarily bypass Wnt/β-catenin asymmetry by downregulating pop-1 expression. Thereby, RNT-1/BRO-1 appears to render POP-1 below the level required for its repressor function, which converts differentiation into self-renewal. Thus, we found that conserved Runx/CBFβ-type stem cell regulators switch asymmetric to proliferative cell division by opposing TCF-related transcriptional repression.

A Tug-of-War between Cell Shape and Polarity Controls Division Orientation to Ensure Robust Patterning in the Mouse Blastocyst.

SummaryOriented cell division patterns tissues by modulating cell position and fate. While cell geometry, junctions, cortical tension, and polarity are known to control division orientation, relatively little is known about how these are coordinated to ensure robust patterning. Here, we systematically characterize cell division, volume, and shape changes during mouse pre-implantation development by in toto live imaging. The analysis leads us to a model in which the apical domain competes with cell shape to determine division orientation. Two key predictions of the model are verified experimentally: when outside cells of the 16-cell embryo are released from cell shape asymmetry, the axis of division is guided by the apical domain. Conversely, orientation cues from the apical domain can be overcome by applied shape asymmetry in the 8-cell embryo. We propose that such interplay between cell shape and polarity in controlling division orientation ensures robust patterning of the blastocyst and possibly other tissues.

From spiral cleavage to bilateral symmetry: the developmental cell lineage of the annelid brain

BackgroundDuring early development, patterns of cell division—embryonic cleavage—accompany the gradual restriction of blastomeres to specific cell fates. In Spiralia, which include annelids, mollusks, and flatworms, “spiral cleavage” produces a highly stereotypic, spiral-like arrangement of blastomeres and swimming trochophore-type larvae with rotational (spiral) symmetry. However, starting at larval stages, spiralian larvae acquire elements of bilateral symmetry, before they metamorphose into fully bilateral juveniles. How this spiral-to-bilateral transition occurs is not known and is especially puzzling for the early differentiating brain and head sensory organs, which emerge directly from the spiral cleavage pattern. Here we present the developmental cell lineage of the Platynereis larval episphere.ResultsLive-imaging recordings from the zygote to the mid-trochophore stage (~ 30 hpf) of the larval episphere of the marine annelid Platynereis dumerilii reveal highly stereotypical development and an invariant cell lineage of early differentiating cell types. The larval brain and head sensory organs develop from 11 pairs of bilateral founders, each giving rise to identical clones on the right and left body sides. Relating the origin of each bilateral founder pair back to the spiral cleavage pattern, we uncover highly divergent origins: while some founder pairs originate from corresponding cells in the spiralian lineage on each body side, others originate from non-corresponding cells, and yet others derive from a single cell within one quadrant. Integrating lineage and gene expression data for several embryonic and larval stages, we find that the conserved head patterning genes otx and six3 are expressed in bilateral founders representing divergent lineage histories and giving rise to early differentiating cholinergic neurons and head sensory organs, respectively.ConclusionsWe present the complete developmental cell lineage of the Platynereis larval episphere, and thus the first comprehensive account of the spiral-to-bilateral transition in a developing spiralian. The bilateral symmetry of the head emerges from pairs of bilateral founders, similar to the trunk; however, the head founders are more numerous and show striking left-right asymmetries in lineage behavior that we relate to differential gene expression.

Towards analytical typologies of plot systems: Quantitative profile of five European cities

The importance of the plot (also referred to as ‘property’) as one of the fundamental elements of urban form is well recognized within the field of urban morphology. Despite the fact that it is often described as the basic element in the pattern of land divisions, which are essential as organizational frameworks for urban form, studies offering comprehensive descriptions and classifications of plot systems are quite scant. The aim of the paper is to introduce a classification of plot systems into typologies based on five European cities, in order to distinguish particular spatial differences and similarities in terms of their plot structure. The proposed typologies are developed using unsupervised k-means cluster analysis based on numeric attributes derived from central theories in urban morphology. The introduced typologies are essentially configurational, allowing collective systematic properties of plot systems to be captured. Numeric attributes include plot differentiation (or plot size), plot frontage and compactness ratio, corresponding to essential qualities of plot systems such as the capacity to carry differences in space, the ability to operate as interface between street and building and providing a framework for evolution of built form over time. All three attributes are translated into configurational measures in order to capture the context of the plot system, rather than the parameters of individual plots. The combination of these deductively defined variables with algorithmically defined classification methods results in seven plot types that can be used to scale up traditional urban morphological analysis to whole city regions and conduct substantial comparison of patterns within, but also between these regions. Further, it also makes it possible to describe commonly recognized plot patterns and discover new ones.

Noncanonical auxin signaling regulates cell division pattern during lateral root development

In both plants and animals, multiple cellular processes must be orchestrated to ensure proper organogenesis. The cell division patterns control the shape of growing organs, yet how they are precisely determined and coordinated is poorly understood. In plants, the distribution of the phytohormone auxin is tightly linked to organogenesis, including lateral root (LR) development. Nevertheless, how auxin regulates cell division pattern during lateral root development remains elusive. Here, we report that auxin activates Mitogen-Activated Protein Kinase (MAPK) signaling via transmembrane kinases (TMKs) to control cell division pattern during lateral root development. Both TMK1/4 and MKK4/5-MPK3/6 pathways are required to properly orient cell divisions, which ultimately determine lateral root development in response to auxin. We show that TMKs directly and specifically interact with and phosphorylate MKK4/5, which is required for auxin to activate MKK4/5-MPK3/6 signaling. Our data suggest that TMK-mediated noncanonical auxin signaling is required to regulate cell division pattern and connect auxin signaling to MAPK signaling, which are both essential for plant development.

Fruit Architecture in Polyamine-Rich Tomato Germplasm Is Determined via a Medley of Cell Cycle, Cell Expansion, and Fruit Shape Genes.

Shape and size are important features of fruits. Studies using tomatoes expressing yeast Spermidine Synthase under either a constitutive or a fruit-ripening promoter showed obovoid fruit phenotype compared to spherical fruit in controls, suggesting that polyamines (PAs) have a role in fruit shape. The obovoid fruit pericarp exhibited decreased cell layers and pericarp thickness compared to wild-type fruit. Transgenic floral buds and ovaries accumulated higher levels of free PAs, with the bound form of PAs being predominant. Transcripts of the fruit shape genes, SUN1 and OVATE, and those of CDKB2, CYCB2, KRP1 and WEE1 genes increased significantly in the transgenic ovaries 2 and 5 days after pollination (DAP). The levels of cell expansion genes CCS52A/B increased at 10 and 20 DAP in the transgenic fruits and exhibited negative correlation with free or bound forms of PAs. In addition, the cell layers and pericarp thickness of the transgenic fruits were inversely associated with free or bound PAs in 10 and 20 DAP transgenic ovaries. Collectively, these results provide evidence for a linkage between PA homeostasis and expression patterns of fruit shape, cell division, and cell expansion genes during early fruit development, and suggest role(s) of PAs in tomato fruit architecture.

MicroRNA Dynamics and Functions During Arabidopsis Embryogenesis.

MicroRNAs (miRNAs) are short noncoding RNAs that mediate the repression of target transcripts in plants and animals. Although miRNAs are required throughout plant development, relatively little is known regarding their embryonic functions. To systematically characterize embryonic miRNAs in Arabidopsis (Arabidopsis thaliana), we developed or applied high-throughput sequencing-based methods to profile hundreds of miRNAs and associated targets throughout embryogenesis. We discovered dozens of miRNAs that dynamically cleave and repress target transcripts, including 30 that encode transcription factors. Transcriptome analyses indicated that these miRNA:target interactions have profound effects on embryonic gene expression programs. Moreover, we demonstrated that the miRNA-mediated repression of six transcription factors are individually required for proper division patterns of various embryonic cell lineages. These data indicate that the miRNA-directed repression of multiple transcription factors is critically important for the establishment of the plant body plan, and they provide a foundation to further investigate how miRNAs contribute to these initial cellular differentiation events.

Live Cell Imaging of Neural Stem Cells in the Drosophila Larval Brain.

Live cell imaging gives valuable insights into the dynamic biological processes within and between cells. An important aspect of live cell imaging is to keep the cells under best physiological condition and to prevent abnormal cellular behavior, which might be caused by phototoxicity during microscopy. In this chapter we describe a protocol to visualize division patterns of neural stem cells in live whole mount brains of Drosophila larvae. We also present a newly developed live cell chamber that allows us to control the environmental air during live cell imaging. The protocol can be adapted to look at a wide range of cellular and tissue behavior in the Drosophila model system.

Nutlin-3-Induced Sensitization of Non-Small Cell Lung Cancer Stem Cells to Axitinib-Induced Apoptosis Through Repression of Akt1/Wnt Signaling.

The aim of this study was to investigate the potential biological activities of nutlin-3 in the regulation of growth and proliferation of non-small cell lung cancer (NSCLC) stem cells (CSCs), which may help in sensitizing to axitinib-induced apoptosis. Nutlin-3 induction of p53 expression was used to test its role in controlling the cell division pattern and apoptosis of NSCLC cells. A549 cells and H460 cells were pretreated with nutlin-3 and then treated with either an Akt1 activator or shRNA-GSK3β, to investigate the potential role of p53 sensitization in the biological effects of axitinib. We also determined the expression levels of GSK3β and p-Akt1 in patients with NSCLC and determined their potential association with survival data using Kaplan–Meier plots and CBIOTAL. Increased p53 expression stimulated the induction of apoptosis by axitinib and promoted asymmetric cell division (ACD) of NSCLC CSCs. The repression of Akt phosphorylation induced by nutlin-3 promoted the ACD of lung CSCs, decreasing the proportion of the stem cell population. In addition to the induction of apoptosis by axitinib through inhibition of Wnt signaling, nutlin-3 treatment further enhanced axitinib-induced apoptosis by inhibiting Akt1/GSK3β/Wnt signaling. The low expression of GSK3β and increased expression of p-Akt in patients with NSCLC were closely associated with the development of NSCLC. TP53 stimulates the induction of apoptosis in NSCLC by axitinib and the ACD of lung CSCs through its regulatory effects on the p53/Akt/GSK3β pathways.

Division pattern of the hypoglossal canal in human fetuses

Amaç: Bu çalışmanın amacı, insan fetüslerinde septum veya çift hipoglossal kanal insidansını tespit etmektir. Yöntem: Tıp Fakültesi Anatomi Anabilim Dalı envanterinde bulunan ortalama 23.36±2.63 haftalık 25 fetüs (14 erkek, 11 kız) çalışmaya dahil edildi. Bulgular: Fetüslerin 9’unda (%36) septum vardı. Fetüslerin 2’sinde (%8) bilateral, 3’ünde (%12) sağ ve 4’ünde (%16) sol tarafta çift hipoglossal kanal tespit edildi. Gestasyonal yaş, cinsiyet ve taraflar ile septum varlığı arasında bir ilişki görülmedi. Sonuç: Hipoglossal kanalın bölümlenme paterni özellikle hipoglossal kanal schwannoma’ları gibi posterior kraniyal fossa operasyonları açısından önem taşıdığından nörocerrahlar ve radyololar için büyük önem taşımaktadır. Ayrıca çalışmamızdaki veriler, kafa tabanının gelişiminin anlaşılması açısından anatomistler ve antropologlar için kıymetli olabilir.

A Toolkit for High Resolution Imaging of Cell Division and Phytohormone Signaling in Legume Roots and Root Nodules.

Legume plants benefit from a nitrogen-fixing symbiosis in association with rhizobia hosted in specialized root nodules. Formation of root nodules is initiated by de novo organogenesis and coordinated infection of these developing lateral root organs by rhizobia. Both bacterial infection and nodule organogenesis involve cell cycle activation and regulation by auxin and cytokinin is tightly integrated in the process. To characterize the hormone dynamics and cell division patterns with cellular resolution during nodulation, sensitive and specific sensors suited for imaging of multicellular tissues are required. Here we report a modular toolkit, optimized in the model legume Lotus japonicus, for use in legume roots and root nodules. This toolkit includes synthetic transcriptional reporters for auxin and cytokinin, auxin accumulation sensors and cell cycle progression markers optimized for fluorescent and bright field microscopy. The developed vectors allow for efficient one-step assembly of multiple units using the GoldenGate cloning system. Applied together with a fluorescence-compatible clearing approach, these reporters improve imaging depth and facilitate fluorescence examination in legume roots. We additionally evaluate the utility of the dynamic gravitropic root response in altering the timing and location of auxin accumulation and nodule emergence. We show that alteration of auxin distribution in roots allows for preferential nodule emergence at the outer side of the bend corresponding to a region of high auxin signaling capacity. The presented tools and procedures open new possibilities for comparative mutant studies and for developing a more comprehensive understanding of legume-rhizobia interactions.

Asymmetric Decentralization and Peace Building: A Comparison of Aceh and Northern Ireland

This article compares the asymmetric form of decentralization model in the Aceh, Indonesia, and in the Northern Ireland, the United Kingdom, towards the process of peacemaking. In general, the decentralization models in the two regions have almost similar backgrounds due to prolonged religious conflicts. However, the devolution model in Northern Ireland is far more complex than Aceh’s, and closely related to other countries, that affect the constitutional status of Northern Ireland in the future. This is different from the Aceh government, which since the Helsinki MoU has remained a part of the Indonesia’s unitary form. On the other hand, in terms of governmental matters, decentralization in Aceh reflects a reduction in their special authority, with a pattern of division of government functions that is almost similar to the division of functions in the ordinary regional government. It is in contrast to the affairs of Northern Ireland’s government concerning matters that are categorized traditionally as the exclusive affairs of the central government in a unitary state, such as security matters. This provides a lesson for Indonesia that the issue of federalism in Aceh is indisputable and the Northern Ireland case reinforces the insight that federal state and unitary state have some degrees of similarity.

Growth and the cell cycle in green algae dividing by multiple fission.

Most cells divide into two daughter cells; however, some green algae can have different division patterns in which a single mother cell can sometimes give rise to up to thousands of daughter cells. Although such cell cycle patterns can be very complex, they are governed by the same general concepts as the most common binary fission. Moreover, cell cycle progression appears to be connected with size, since cells need to ensure that their size after division will not drop below the limit required for survival. Although the exact mechanism that lets cells measure cell size remains largely unknown, there have been several prominent hypotheses that try to explain it.

Role of PRY-1/Axin in heterochronic miRNA-mediated seam cell development

BackgroundCaenorhabditis elegans seam cells serve as a good model to understand how genes and signaling pathways interact to control asymmetric cell fates. The stage-specific pattern of seam cell division is coordinated by a genetic network that includes WNT asymmetry pathway components WRM-1, LIT-1, and POP-1, as well as heterochronic microRNAs (miRNAs) and their downstream targets. Mutations in pry-1, a negative regulator of WNT signaling that belongs to the Axin family, were shown to cause seam cell defects; however, the mechanism of PRY-1 action and its interactions with miRNAs remain unclear.ResultsWe found that pry-1 mutants in C. elegans exhibit seam cell, cuticle, and alae defects. To examine this further, a miRNA transcriptome analysis was carried out, which showed that let-7 (miR-48, miR-84, miR-241) and lin-4 (lin-4, miR-237) family members were upregulated in the absence of pry-1 function. Similar phenotypes and patterns of miRNA overexpression were also observed in C. briggsae pry-1 mutants, a species that is closely related to C. elegans. RNA interference-mediated silencing of wrm-1 and lit-1 in the C. elegans pry-1 mutants rescued the seam cell defect, whereas pop-1 silencing enhanced the phenotype, suggesting that all three proteins are likely important for PRY-1 function in seam cells. We also found that these miRNAs were overexpressed in pop-1 hypomorphic animals, suggesting that PRY-1 may be required for POP-1-mediated miRNA suppression. Analysis of the let-7 and lin-4-family heterochronic targets, lin-28 and hbl-1, showed that both genes were significantly downregulated in pry-1 mutants, and furthermore, lin-28 silencing reduced the number of seam cells in mutant animals.ConclusionsOur results show that PRY-1 plays a conserved role to maintain normal expression of heterochronic miRNAs in nematodes. Furthermore, we demonstrated that PRY-1 acts upstream of the WNT asymmetry pathway components WRM-1, LIT-1, and POP-1, and miRNA target genes in seam cell development.

A Rule of Law Approach to the Division of Authority of Office in China

Making the authority of office of government at all levels more standardized and law-based is an important step in the process of deepening financial reform and establishing fiscal and taxation systems governed by law. To do so, China needs to clarify further the legal division of authority of government at every level and protect and promote its exercise through legal mechanisms. Governments are given powers so that they may effectively protect citizens’ basic interests. On the premise of clarifying the boundaries of government functions, we should use legal means and observe the principle of restraint in determining the scope of authority of office. The criteria for the division have both economic and legal aspects. Economic criteria are more basic; legal criteria make adjustments to the pattern of division derived from economic criteria. The authority of office can be divided into legislative supervision and enforcement. The current legal system allocates powers of legislative supervision quite rationally, but supervision of judicial powers in particular should be appropriately centralized. In terms of enforcement, some space is left for the division of administrative powers in the current system, and there is quite obvious homogeneity of powers between different levels of government. The legal division of authority of office requires that basic principles be stipulated in the Constitution and that a basic fiscal law be devised that clarifies the powers of all levels of government. Allocation of fiscal powers between governments and optimization of the transfer payment system, together with budget constraints, will provide an institutional guarantee for the division of these powers. Fundamentally, the legal confirmation of a system of civil rights can promote the implementation of statutory powers.

The cleavage pattern of calanoid copepods-a case study.

Many crustacean groups show stereotyped cleavage patterns during early ontogeny. However, these patterns differ between the various major crustacean taxa, and a general mode is difficult to extract. Previous studies suggested that also copepods undergo an early cleavage with a more or less stereotyped pattern of blastomere divisions and fates. Yet, copepod embryology has been largely neglected. The last investigation of this kind dates back more than a century and the results are somewhat contradictory when compared with those of other researchers. To overcome these problems, we studied the early development of a so far undescribed calanoid copepod species, Skistodiaptomus sp., applying histochemical staining, confocal laser scanning microscopy, and bifocal 4D microscopy. The blastomere arrangement of the four-cell stage of this species varies to a large degree. It can either form a typical radial pattern with the four blastomeres lying in one plane or a tilted orientation of the axes connecting the sister cells of the previous division. In both cases, a stereotyped division pattern is maintained inside each quadrant during subsequent cleavages. In addition, we found two types of blastomere arrangements with a mirror symmetry. Most divisions within the quadrants follow the perpendicularity rule until the eighth cleavage. Deviations from this rule occur only in the narrow regions where the different quadrants touch and near the site of gastrulation. Gastrulation is initiated around the descendants of one individually identifiable blastomere of the 16-cell stage. This cell divides in a specific manner forming a characteristic cell arrangement, the gastrulation triangle. This gastrulation triangle initiates the internalization process of the gastrulation and it is encircled by another characteristic cell type, the crown cells. Our observations reveal several similarities to the early development of Calanus finmarchicus, another calanoid species. These relate to blastomere arrangements and divisions and the pattern of gastrulation. As Calanoida represent a basal or near basal branch of the copepod tree, this description will provide the ground for reconstruction of the cleavage pattern of the last common ancestor of Copepoda.