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<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" article-type="research-article" dtd-version="1.2" xml:lang="en"><front><journal-meta><journal-id journal-id-type="publisher-id">Morphology</journal-id><journal-title-group><journal-title xml:lang="en">Morphology</journal-title><trans-title-group xml:lang="ru"><trans-title>Морфология</trans-title></trans-title-group></journal-title-group><issn publication-format="print">1026-3543</issn><issn publication-format="electronic">2949-2556</issn><publisher><publisher-name xml:lang="en">Eco-Vector</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="publisher-id">688247</article-id><article-id pub-id-type="doi">10.17816/morph.688247</article-id><article-id pub-id-type="edn">YTEHPS</article-id><article-categories><subj-group subj-group-type="toc-heading" xml:lang="en"><subject>Original Study Articles</subject></subj-group><subj-group subj-group-type="toc-heading" xml:lang="ru"><subject>Оригинальные исследования</subject></subj-group><subj-group subj-group-type="article-type"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title xml:lang="en">Analysis of the structural organization of the metatarsal bones and toe phalanges of the human foot using anatomical network models</article-title><trans-title-group xml:lang="ru"><trans-title>Анализ структурной организации костей плюсны и фаланг пальцев стопы человека по данным анатомических сетевых моделей</trans-title></trans-title-group><trans-title-group xml:lang="zh"><trans-title>基于解剖网络模型数据的人类足部跖骨和趾骨结构组织分析</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-6836-5650</contrib-id><contrib-id contrib-id-type="spin">1898-3355</contrib-id><name-alternatives><name xml:lang="en"><surname>Gayvoronsky</surname><given-names>Ivan V.</given-names></name><name xml:lang="ru"><surname>Гайворонский</surname><given-names>Иван Васильевич</given-names></name><name xml:lang="zh"><surname>Gayvoronsky</surname><given-names>Ivan V.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>MD, Dr. Sci. (Medicine), Professor</p></bio><bio xml:lang="ru"><p>д-р мед. наук, профессор</p></bio><bio xml:lang="zh"><p>MD, Dr. Sci. (Medicine), Professor</p></bio><email>i.v.gaivoronsky@mail.ru</email><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-3857-7388</contrib-id><contrib-id contrib-id-type="spin">8320-9471</contrib-id><name-alternatives><name xml:lang="en"><surname>Fandeeva</surname><given-names>Oksana M.</given-names></name><name xml:lang="ru"><surname>Фандеева</surname><given-names>Оксана Маликовна</given-names></name><name xml:lang="zh"><surname>Fandeeva</surname><given-names>Oksana M.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>MD,<bold> </bold>Dr. Sci. (Medicine)</p></bio><bio xml:lang="ru"><p>д-р мед. наук</p></bio><bio xml:lang="zh"><p>MD,<bold> </bold>Dr. Sci. (Medicine)</p></bio><email>osteolog_oxana@mail.ru</email><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff3"/></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0007-4987-9262</contrib-id><contrib-id contrib-id-type="spin">1035-3261</contrib-id><name-alternatives><name xml:lang="en"><surname>Pashchenko</surname><given-names>Pavel S.</given-names></name><name xml:lang="ru"><surname>Пащенко</surname><given-names>Павел Степанович</given-names></name><name xml:lang="zh"><surname>Pashchenko</surname><given-names>Pavel S.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>MD, Dr. Sci. (Medicine), Professor</p></bio><bio xml:lang="ru"><p>д-р мед. наук, профессор</p></bio><bio xml:lang="zh"><p>MD, Dr. Sci. (Medicine), Professor</p></bio><email>pashchenko@vmeda.org</email><xref ref-type="aff" rid="aff1"/></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-5569-7325</contrib-id><contrib-id contrib-id-type="spin">3532-1203</contrib-id><name-alternatives><name xml:lang="en"><surname>Nichiporuk</surname><given-names>Gennady I.</given-names></name><name xml:lang="ru"><surname>Ничипорук</surname><given-names>Геннадий Иванович</given-names></name><name xml:lang="zh"><surname>Nichiporuk</surname><given-names>Gennady I.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>MD, Cand. Sci. (Medicine), Assistant Professor</p></bio><bio xml:lang="ru"><p>канд. мед. наук, доцент</p></bio><bio xml:lang="zh"><p>MD, Cand. Sci. (Medicine), Assistant Professor</p></bio><email>nichiporuk@vmeda.org</email><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-8429-5840</contrib-id><contrib-id contrib-id-type="spin">9343-0818</contrib-id><name-alternatives><name xml:lang="en"><surname>Vinogradov</surname><given-names>Stas V.</given-names></name><name xml:lang="ru"><surname>Виноградов</surname><given-names>Стас Викторович</given-names></name><name xml:lang="zh"><surname>Vinogradov</surname><given-names>Stas V.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>MD, Cand. Sci. (Medicine), Assistant Professor</p></bio><bio xml:lang="ru"><p>канд. мед. наук, доцент</p></bio><bio xml:lang="zh"><p>MD, Cand. Sci. (Medicine), Assistant Professor</p></bio><email>vitista@mail.ru</email><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-7360-5970</contrib-id><contrib-id contrib-id-type="spin">6861-3767</contrib-id><name-alternatives><name xml:lang="en"><surname>Shashkov</surname><given-names>Vladimir A.</given-names></name><name xml:lang="ru"><surname>Шашков</surname><given-names>Владимир Андреевич</given-names></name><name xml:lang="zh"><surname>Shashkov</surname><given-names>Vladimir A.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>MD, Cand. Sci. (Medicine)</p></bio><bio xml:lang="ru"><p>канд. мед. наук</p></bio><bio xml:lang="zh"><p>MD, Cand. Sci. (Medicine)</p></bio><email>dr.shashkov@mail.ru</email><xref ref-type="aff" rid="aff1"/></contrib></contrib-group><aff-alternatives id="aff1"><aff><institution xml:lang="en">Kirov Military Medical Academy</institution></aff><aff><institution xml:lang="ru">Военно-медицинская академия им. С.М. Кирова</institution></aff><aff><institution xml:lang="zh">Kirov Military Medical Academy</institution></aff></aff-alternatives><aff-alternatives id="aff2"><aff><institution xml:lang="en">Saint-Petersburg State University</institution></aff><aff><institution xml:lang="ru">Санкт-Петербургский государственный университет</institution></aff><aff><institution xml:lang="zh">Saint-Petersburg State University</institution></aff></aff-alternatives><aff-alternatives id="aff3"><aff><institution xml:lang="en">Academician I.P. Pavlov First St. Petersburg State Medical University</institution></aff><aff><institution xml:lang="ru">Первый Санкт-Петербургский государственный медицинский университет им. акад. И.П. Павлова</institution></aff><aff><institution xml:lang="zh">Academician I.P. Pavlov First St. Petersburg State Medical University</institution></aff></aff-alternatives><pub-date date-type="preprint" iso-8601-date="2026-03-01" publication-format="electronic"><day>01</day><month>03</month><year>2026</year></pub-date><pub-date date-type="pub" iso-8601-date="2026-08-07" publication-format="electronic"><day>07</day><month>08</month><year>2026</year></pub-date><volume>164</volume><issue>3</issue><issue-title xml:lang="en"/><issue-title xml:lang="ru"/><issue-title xml:lang="zh"/><fpage>315</fpage><lpage>325</lpage><history><date date-type="received" iso-8601-date="2025-07-24"><day>24</day><month>07</month><year>2025</year></date><date date-type="accepted" iso-8601-date="2025-11-11"><day>11</day><month>11</month><year>2025</year></date></history><permissions><copyright-statement xml:lang="en">Copyright ©; 2026, Eco-Vector</copyright-statement><copyright-statement xml:lang="ru">Copyright ©; 2026, Эко-Вектор</copyright-statement><copyright-statement xml:lang="zh">Copyright ©; 2026,</copyright-statement><copyright-year>2026</copyright-year><copyright-holder xml:lang="en">Eco-Vector</copyright-holder><copyright-holder xml:lang="ru">Эко-Вектор</copyright-holder><ali:free_to_read xmlns:ali="http://www.niso.org/schemas/ali/1.0/" start_date="2029-08-07"/><license><ali:license_ref xmlns:ali="http://www.niso.org/schemas/ali/1.0/">https://creativecommons.org/licenses/by-nc-nd/4.0/</ali:license_ref></license></permissions><self-uri xlink:href="https://j-morphology.com/1026-3543/article/view/688247">https://j-morphology.com/1026-3543/article/view/688247</self-uri><abstract xml:lang="en"><p><bold>BACKGROUND: </bold>The human forefoot, despite its fundamental role in bipedalism, is the least studied part of the foot in most biomechanical and clinical models, where it is often considered a rigid structure. Evolutionary adaptations associated with upright walking have led to a complex morphological structure of the forefoot. Existing concepts of morphological modularity and integration are often limited by the complexity of analyzing multiple relationships. This study suggests solving this problem by applying anatomical network analysis (AnNA) to study the structural organization of the forefoot. AnNA enables formalization of the connections between the components of the foot, revealing patterns of its integration and modularity.</p> <p><bold>AIM: </bold>To study the structural organization of the metatarsal bones and toe phalanges of the human foot using AnNA.</p> <p><bold>METHODS:</bold> As part of this study, the igraph library in the R programming environment was used to analyze the AnNA network. Modeling and visualization of the network structure were carried out using the Fruchterman–Reingold algorithm, which ensures the optimal placement of nodes. Structural characteristics, including modularity and integration of the AnNA network, were analyzed using the Spinglass algorithm. Osteometric parameters — the length of the metatarsal bones and phalanges of toes I–V of the right foot from 100 middle-aged men — were used as input data for the AnNA analysis.</p> <p><bold>RESULTS:</bold> The structural organization of the metatarsal bones and toe phalanges of the human foot consists of several modules: a proximal module (includes metatarsals I–V) and a distal module (includes the phalanges of toes I–V) relative to the longitudinal axis of the foot, and a medial module (includes metatarsals I–III and the phalanges of toes I–III) and a lateral module (includes metatarsals IV–V and the phalanges of toes IV–V) relative to the transverse axis of the foot. Network analysis showed that the distal and medial modules are more complex and less anisomeric than the proximal and lateral modules. The distal module is more complex than the proximal one, while the medial and lateral modules show greater similarity in organizational complexity. Overall, the medial and lateral modules are more similar to each other than are the distal and proximal modules.</p> <p><bold>CONCLUSION: </bold>Morphological modularity and integration are organizing principles of the bone architecture of the foot. The structural organization of the metatarsal bones and toe phalanges of the human foot is a system of four modules — a proximal (metatarsal bones) and a distal (toe phalanges) module relative to the longitudinal axis of the foot, and a medial (metatarsals I–III and phalanges of toes I–III) and a lateral (metatarsals IV–V and phalanges of toes IV–V) module relative to the transverse axis of the foot.</p></abstract><trans-abstract xml:lang="ru"><p><bold>Обоснование. </bold>Передний отдел стопы человека, несмотря на его фундаментальную роль в бипедализме, является наименее изученной частью, поскольку в большинстве биомеханических и клинических моделей походки он рассматривается как жёсткая конструкция. Эволюционные адаптации, связанные с прямохождением, обусловили формирование сложной морфологической структуры переднего отдела стопы. Существующие представления о морфологической модульности и интеграции часто ограничены сложностью анализа множественных взаимосвязей. Данное исследование предлагает решить эту проблему, применяя анатомический сетевой анализ (Anatomical Network Analysis, AnNA) для изучения структурной организации переднего отдела стопы. AnNA позволяет формализовать связи между компонентами стопы, выявляя закономерности её интеграции и модульности.</p> <p><bold>Цель исследования</bold> — изучить структурную организацию костей плюсны и фаланг пальцев стопы человека методом анатомического сетевого анализа.</p> <p><bold>Методы.</bold> В рамках данного исследования для анализа сети AnNA применена библиотека <italic>igraph</italic> в среде программирования R. Моделирование и визуализация структуры сети выполнены с использованием алгоритма Fruchterman–Reingold, обеспечивающего оптимальное расположение узлов. Анализ структурных характеристик, включая модульность и интеграцию сети AnNA, проводили с использованием алгоритма Spinglass. В качестве входных данных для AnNA использованы остеометрические параметры ста мужчин среднего возраст — длина плюсневых костей и фаланг I–V пальцев правой стопы.</p> <p><bold>Результаты. </bold>Структурная организация костей плюсны и фаланг пальцев стопы представлена несколькими модулями: относительно продольной оси стопы — проксимальным (включает I–V плюсневые кости) и дистальным (включает фаланги I–V пальцев); относительно поперечной оси стопы — медиальным (включает I–III плюсневые кости и фаланги I–III пальцев) и латеральным (включает IV–V плюсневые кости и фаланги IV–V пальцев). Сетевой анализ показал, что дистальный и медиальный модули являются более сложными и менее анизомерными, чем проксимальный и латеральный. Дистальный модуль организован более сложно, чем проксимальный, а медиальный и латеральный модули демонстрируют более высокую схожесть в сложности организации. В целом медиальный и латеральный модули более сходны между собой, чем дистальный и проксимальный.</p> <p><bold>Заключение.</bold> Морфологическая модульность и интеграция являются организующими факторами костной архитектуры стопы. Структурная организация костей плюсны и фаланг пальцев стопы представляет собой систему, состоящую из четырёх модулей — проксимального (плюсневые кости) и дистального (фаланги пальцев) относительно продольной оси стопы, медиального (I–III плюсневые кости и фаланги I–III пальцев) и латерального (IV–V плюсневые кости и фаланги IV–V пальцев) относительно поперечной оси.</p></trans-abstract><trans-abstract xml:lang="zh"><p><bold>论证。</bold>尽管人类前足在双足行走中起着至关重要的作用，但在大多数生物力学和临床模型中，它却是足部研究最少的部分，通常被视为一个刚性结构。与直立行走相关的进化适应导致了前足复杂的形态结构。现有的形态模块化和整合概念常常受限于分析多种关系的复杂性。本研究旨在通过应用解剖网络分析 (AnNA) 来研究前足的结构组织，从而解决这一问题。AnNA 能够形式化地描述足部各组成部分之间的关系，揭示其整合和模块化的模式。</p> <p><bold>目的。</bold>本研究旨在利用解剖网络分析方法研究人类足部跖骨和趾骨的结构组织。</p> <p><bold>方法。</bold>本研究使用 R 编程环境中的 igraph 库来分析 AnNA (解剖网络分析) 网络。网络结构的建模和可视化采用 Fruchterman-Reingold 算法，该算法能够确保节点的最佳放置。利用 Spinglass 算法对 AnNA 网络的结构特征（包括模块化和整合度）进行了分析。AnNA 的输入数据为 100 名中年男性的骨骼测量参数，即右脚第一至第五趾的跖骨和趾骨长度。</p> <p><bold>结果。</bold>足部跖骨和趾骨的结构组织由多个模块构成：相对于足纵轴，可分为近端模块（包括第 1-5 跖骨）和远端模块（包括第 1-5 趾骨）；相对于足横轴，可分为内侧模块（包括第 1-3 跖骨和第 1-3 趾骨）和外侧模块（包括第 4-5 跖骨和第 4-5 趾骨）。网络分析表明，远端模块和内侧模块比近端模块和外侧模块更复杂，且异尺性程度更低。远端模块的组织方式比近端模块更复杂，而内侧模块和外侧模块在组织复杂性方面表现出更高的相似性。总体而言，内侧模块和外侧模块之间的相似性高于远端模块和近端模块之间的相似性。</p> <p><bold>结论。</bold>形态模块化和整合是足部骨骼结构组织化的关键因素。跖骨和趾骨的结构组织是一个由四个模块组成的系统：相对于足纵轴，分为近端模块（跖骨）和远端模块（趾骨）；相对于横轴，分为内侧模块（第 1-3 跖骨和第 1-3 趾的趾骨）和外侧模块（第 4-5 跖骨和第 4-5 趾的趾骨）。</p></trans-abstract><kwd-group xml:lang="en"><kwd>human foot</kwd><kwd>forefoot</kwd><kwd>structural organization</kwd><kwd>anatomical network analysis</kwd></kwd-group><kwd-group xml:lang="ru"><kwd>стопа человека</kwd><kwd>передний отдел стопы</kwd><kwd>структурная организация</kwd><kwd>анатомический сетевой анализ</kwd></kwd-group><kwd-group xml:lang="zh"><kwd>人足</kwd><kwd>前足</kwd><kwd>结构组织</kwd><kwd>解剖网络分析</kwd></kwd-group><funding-group/></article-meta></front><body></body><back><ref-list><ref id="B1"><label>1.</label><mixed-citation>Mahieu C, Salvia P, Beyer B, et al. 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