Novel Mitochondria-Targeted Heat-Soluble Proteins Identified in the Anhydrobiotic Tardigrade Improve Osmotic Tolerance of Human Cells

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Fukuda, Y., & Tsuyoshi Inoue, A. (2018) · Protein Science · doi:10.1002/pro.3393 · Cited by 21 · see citing works

Towards water-free biobanks: long-term dry-preservation at room temperature of desiccation-sensitive enzyme luciferase in air-dried insect cells

Kikuta, S., Watanabe, S. J., Sato, R., Gusev, O., Nesmelov, A. A., Sogame, Y., Cornette, R., & Kikawada, T. (2017) · Scientific Reports · doi:10.1038/s41598-017-06945-y · Cited by 21 · see citing works

Small LEA proteins mitigate air-water interface damage to fragile cryo-EM samples during plunge freezing

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The tardigrade cuticle

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Tardigrade secretory proteins protect biological structures from desiccation

Lim, S., Reilly, C. B., Barghouti, Z., Marelli, B., Way, J. C., & Silver, P. A. (2024) · Communications Biology · doi:10.1038/s42003-024-06336-w · Cited by 18 · see citing works

Structural study of the intrinsically disordered tardigrade damage suppressor protein (Dsup) and its complex with DNA

Zarubin, M. P., Murugova, T. N., Ryzhykau, Y. L., Ivankov, O. I., Uversky, V. N., & Kravchenko, E. (2024) · Scientific Reports · doi:10.1038/s41598-024-74335-2 · Cited by 18 · see citing works

Antioxidant Defense in the Toughest Animals on the Earth: Its Contribution to the Extreme Resistance of Tardigrades

Sadowska‐Bartosz, I., & Bartosz, G. (2024) · International Journal of Molecular Sciences · doi:10.3390/ijms25158393 · Cited by 16 · see citing works

Tardigrade proteins: molecular tools in the phenomenon of anhydrobiosis

Krakowiak, M., Bartylak, T., Kmita, H., Kaczmarek, Ł., & Nawrot, R. (2023) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlad066 · Cited by 15 · see citing works

High quality genome assembly of the anhydrobiotic midge provides insights on a single chromosome-based emergence of extreme desiccation tolerance

Yoshida, Y., Shaikhutdinov, N., Kozlova, O., Itoh, M., Tagami, M., Murata, M., Nishiyori-Sueki, H., Kojima-Ishiyama, M., Noma, S., Cherkasov, A. V., et al. (2022) · NAR Genomics and Bioinformatics · doi:10.1093/nargab/lqac029 · Cited by 15 · see citing works

AMPK activity is required for the induction of anhydrobiosis in a tardigrade Hypsibius exemplaris, and its potential up‐regulator is PP2A

Kondo, K., Mori, M., Tomita, M., & Arakawa, K. (2019) · Genes to Cells · doi:10.1111/gtc.12726 · Cited by 14 · see citing works

Applicable Life-History and Molecular Traits for Studying the Effects of Anhydrobiosis on Aging in Tardigrades

Nagwani, A. K., Kaczmarek, Ł., & Kmita, H. (2022) · Diversity · doi:10.3390/d14080664 · Cited by 13 · see citing works

Increasing temperature-driven changes in life history traits and gene expression of an Antarctic tardigrade species

Giovannini, I., Manfrin, C., Greco, S., Vincenzi, J., Altiero, T., Guidetti, R., Giulianini, P. G., & Rebecchi, L. (2023) · Frontiers in Physiology · doi:10.3389/fphys.2023.1258932 · Cited by 13 · see citing works

Dynamic conformational changes of a tardigrade group-3 late embryogenesis abundant protein modulate membrane biophysical properties

Li, X.-H., Yu, C. W. H., Gomez-Navarro, N., Stancheva, V., Zhu, H., Murthy, A., Wozny, M., Malhotra, K., Johnson, C. M., Blackledge, M., et al. (2023) · PNAS Nexus · doi:10.1093/pnasnexus/pgae006 · Cited by 11 · see citing works

Nondomain biopolymers: Flexible molecular strategies to acquire biological functions

Arakawa, K., Hirose, T., Inada, T., Ito, T., Kai, T., Oyama, M., Tomari, Y., Yoda, T., & Nakagawa, S. (2023) · Genes to Cells · doi:10.1111/gtc.13050 · Cited by 11 · see citing works

Helicity of a tardigrade disordered protein contributes to its protective function during desiccation

Biswas, S., Gollub, E., Yu, F., Ginell, G. M., Holehouse, A. S., Sukenik, S., & Boothby, T. C. (2023) · Protein Science · doi:10.1002/pro.4872 · Cited by 11 · see citing works

Chemobiosis reveals tardigrade tun formation is dependent on reversible cysteine oxidation

Smythers, A. L., Joseph, K. M., O’Dell, H. M., Clark, T. A., Crislip, J. R., Flinn, B. B., Daughtridge, M. H., Stair, E. R., Mubarek, S. N., Lewis, H. C., et al. (2024) · PLoS ONE · doi:10.1371/journal.pone.0295062 · Cited by 11 · see citing works

Toxicity with Waste-Generated Ionizing Radiations: Blunders Behind the Scenes

Sharma, A., Chand, K., Singh, G. B., & Mudgal, G. (2021) · Molecular and integrative toxicology · doi:10.1007/978-3-030-83446-3_15 · Cited by 10 · see citing works

Deciphering the Biological Enigma—Genomic Evolution Underlying Anhydrobiosis in the Phylum Tardigrada and the Chironomid Polypedilum vanderplanki

Yoshida, Y., & Tanaka, S. (2022) · Insects · doi:10.3390/insects13060557 · Cited by 10 · see citing works

Crystal structure of Kumaglobin: a hexacoordinated heme protein from an anhydrobiotic tardigrade, Ramazzottius varieornatus

Kim, J., Fukuda, Y., & Tsuyoshi Inoue, A. (2018) · FEBS Journal · doi:10.1111/febs.14713 · Cited by 10 · see citing works

The wealth of shared resources: Improving molecular taxonomy using eDNA and public databases

Fleming, J. F. (2023) · Zoologica Scripta · doi:10.1111/zsc.12591 · Cited by 10 · see citing works

Acoustomicrofluidic separation of tardigrades from raw cultures for sample preparation

Afzal, M., Park, J., Destgeer, G., Ahmed, H., Iqrar, S. A., Kim, S., Kang, S., Alazzam, A., Yoon, T., & Sung, H. J. (2019) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlz079 · Cited by 10 · see citing works

Elevated external temperature affects cell ultrastructure and heat shock proteins (HSPs) in Paramacrobiotus experimentalis Kaczmarek, Mioduchowska, Poprawa, & Roszkowska, 2020

Kayastha, P., Wieczorkiewicz, F., Pujol, M., Robinson, A., Michalak, M., Kaczmarek, Ł., & Poprawa, I. (2024) · Scientific Reports · doi:10.1038/s41598-024-55295-z · Cited by 9 · see citing works

Sexual dimorphism in the tardigrade Paramacrobiotus metropolitanus transcriptome

Sugiura, K., Yoshida, Y., Hayashi, K., Arakawa, K., Kunieda, T., & Matsumoto, M. (2024) · Zoological Letters · doi:10.1186/s40851-024-00233-0 · Cited by 9 · see citing works

Horizontal Gene Transfer in Metazoa: Examples and Methods

Yoshida, Y., Nowell, R. W., Arakawa, K., & Blaxter, M. (2019) · Springer eBooks · doi:10.1007/978-3-030-21862-1_7 · Cited by 9 · see citing works

Protective roles of highly conserved motif 1 in tardigrade cytosolic‐abundant heat soluble protein in extreme environments

Kang, D., Yang, M. J., Kim, H., & Park, C. (2024) · Protein Science · doi:10.1002/pro.4913 · Cited by 8 · see citing works

Transcriptome analysis of the tardigrade Hypsibius exemplaris exposed to the DNA-damaging agent bleomycin

Yoshida, Y., Hirayama, A., & Arakawa, K. (2024) · Proceedings of the Japan Academy Series B · doi:10.2183/pjab.pjab.100.023 · Cited by 8 · see citing works

Molecular Biology in Tardigrades

Boothby, T. C. (2018) · Zoological monographs · doi:10.1007/978-3-319-95702-9_13 · Cited by 8 · see citing works

The tardigrade-derived mitochondrial abundant heat soluble protein improves adipose-derived stem cell survival against representative stressors

Rolsma, J. L., Darch, W., Higgins, N. C., & Morgan, J. T. (2024) · Scientific Reports · doi:10.1038/s41598-024-62693-w · Cited by 7 · see citing works

An evaluation of thermal tolerance in six tardigrade species in an active and dry state

Loeffelholz, J., Meese, E. K., Giovannini, I., Ullibarri, K., Momeni, S., Merfeld, N., Wessel, J., Guidetti, R., Rebecchi, L., & Boothby, T. C. (2024) · Biology Open · doi:10.1242/bio.060485 · Cited by 7 · see citing works

The major inducible small heat shock protein HSP20-3 in the tardigrade Ramazzottius varieornatus forms filament-like structures and is an active chaperone

Al-Ansari, M., Fitzsimons, T., Wei, W., Goldberg, M. W., Kunieda, T., & Quinlan, R. A. (2024) · Cell Stress and Chaperones · doi:10.1016/j.cstres.2023.12.001 · Cited by 7 · see citing works

Osmotic stress tolerance and transcriptomic response of Ramazzottius varieornatus (Eutardigrada: Ramazzottiidae) following tun formation

Emdee, N., Møbjerg, A., Grollmann, M. M., & Møbjerg, N. (2023) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlad046 · Cited by 7 · see citing works

Tolerance against exposure to solution of magnesium perchlorate in microinvertebrates

Kayastha, P., Rzymski, P., Gołdyn, B., Nagwani, A. K., Fiałkowska, E., Pajdak‐Stós, A., Sobkowiak, R., Robotnikowski, G., & Kaczmarek, Ł. (2023) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlad060 · Cited by 7 · see citing works

Metabolite phosphatase from anhydrobiotic tardigrades

Kato, S., Deguchi, K., Obana, M., Fujio, Y., Fukuda, Y., & Tsuyoshi Inoue, A. (2024) · FEBS Journal · doi:10.1111/febs.17296 · Cited by 6 · see citing works

Tardigrade inspired polyelectrolyte complexation and functional materials

Zhang, F., Song, C., Zhu, X., Gong, J., & Zhao, Q. (2019) · Journal of Materials Chemistry A · doi:10.1039/c9ta10931e · Cited by 6 · see citing works

Tardigrada: An Emerging Animal Model to Study the Endoplasmic Reticulum Stress Response to Environmental Extremes

Kaczmarek, Ł. (2021) · Progress in molecular and subcellular biology · doi:10.1007/978-3-030-67696-4_14 · Cited by 6 · see citing works

A bacterial expression cloning screen reveals single-stranded DNA-binding proteins as potent desicco-protectants

Hibshman, J. D., Clark-Hachtel, C. M., Bloom, K. S., & Goldstein, B. (2024) · Cell Reports · doi:10.1016/j.celrep.2024.114956 · Cited by 5 · see citing works

Protein Intrinsic Disorder and Adaptation to Extreme Environments: Resilience of Chaos

Uversky, V. N. (2025) · Journal of Molecular Biology · doi:10.1016/j.jmb.2025.169547 · Cited by 5 · see citing works

Tardigrade Research in Japan

Suzuki, A. (2016) · Diversity and commonality in animals · doi:10.1007/978-4-431-56432-4_10 · Cited by 5 · see citing works

Crystal structure of hexacoordinated heme protein from anhydrobiotic tardigrade at pH 7

Kim, J., Fukuda, Y., & Tsuyoshi Inoue, A. (2018) · Worldwide Protein Data Bank · doi:10.2210/pdb5zm9/pdb · Cited by 5 · see citing works

A novel Mn-dependent peroxidase contributes to tardigrade anhydrobiosisPreprint

Yoshida, Y., Satoh, T., Ota, C., Tanaka, S., Horikawa, D. D., Tomita, M., Kato, K., & Arakawa, K. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.11.06.370643 · Cited by 5 · see citing works

A comparative ultrastructure study of the tardigrade Ramazzottius varieornatus in the hydrated state, after desiccation and during the process of rehydration

Galas, S., Goff, E. L., Cazevieille, C., Tanaka, A., Cuq, P., Baghdiguian, S., Kunieda, T., Godefroy, N., & Richaud, M. (2024) · PLoS ONE · doi:10.1371/journal.pone.0302552 · Cited by 4 · see citing works

Expressing intrinsically-disordered tardigrade proteins has positive effects on acute but not chronic stress tolerance in Saccharomyces cerevisiae

López, M. I. S., Wheeldon, I. R., & Morgan, J. T. (2025) · PLoS ONE · doi:10.1371/journal.pone.0325682 · Cited by 4 · see citing works

Surviving desiccation: key factors underlying tolerance in prokaryotes and eukaryotes

Rosete-Enríquez, M., Juárez-González, V. R., Escobar-Muciño, E., Muñoz‐Rojas, J., & Quintero‐Hernández, V. (2025) · PROTOPLASMA · doi:10.1007/s00709-025-02134-1 · Cited by 4 · see citing works

Possible roles of CAHS proteins from Tardigrade in osmotic stress tolerance in mammalian cells

Bino, T., Goto, Y., Maryu, G., Arakawa, K., & Aoki, K. (2024) · Cell Structure and Function · doi:10.1247/csf.24035 · Cited by 4 · see citing works

Expression of tardigrade disordered proteins impacts the tolerance to biofuels in a model cyanobacterium Synechocystis sp. PCC 6803

Zhang, H., Liu, Q., Liang, Q., Wang, B., Chen, Z., & Wang, J. (2023) · Frontiers in Microbiology · doi:10.3389/fmicb.2022.1091502 · Cited by 3 · see citing works

Osmolyte-IDP interactions during desiccation

Nicholson, V. M., Meese, E. K., & Boothby, T. C. (2024) · Progress in molecular biology and translational science · doi:10.1016/bs.pmbts.2024.08.007 · Cited by 2 · see citing works

No structure, no problem: Protein stabilization by Hero proteins and other chaperone-like IDPs

Lam, A. Y. W., Tomari, Y., & Tsuboyama, K. (2025) · Biochimica et Biophysica Acta (BBA) - General Subjects · doi:10.1016/j.bbagen.2025.130786 · Cited by 2 · see citing works

Fast autooxidation of a bis-histidyl-ligated globin from the anhydrobiotic tardigrade, Ramazzottius varieornatus, by molecular oxygen

Kobayashi, K., Kim, J., Fukuda, Y., Kozawa, T., & Tsuyoshi Inoue, A. (2021) · The Journal of Biochemistry · doi:10.1093/jb/mvab003 · Cited by 2 · see citing works

Comparative transcriptomics reveal a novel tardigrade specific DNA binding protein induced in response to ionizing radiationPreprint

Anoud, M., Delagoutte, E., Helleu, Q., Brion, A., Duvernois‐Berthet, E., As, M., Marques, X., Lamribet, K., Sénamaud‐Beaufort, C., Jourdren, L., et al. (2024) · eLife · doi:10.7554/elife.92621.2 · Cited by 2 · see citing works

Mitochondria-Derived Reactive Oxygen Species Regulation of Tardigrade Osmobiosis Revealed by Proteomics of Hypsibius exemplaris

Stair, E. R., Flinn, B. B., Grimmett, A., Balboa, S. J., Burr, A. J., Siebold, J. A., Baca, A. G., Daughtridge, M. H., Kolling, D. R. J., & Hicks, L. M. (2025) · Journal of Proteome Research · doi:10.1021/acs.jproteome.5c00222 · Cited by 1 · see citing works

Expression of tardigrade extremotolerance-associated proteins improves recovery but not acute stress response of human microvascular endothelial cells

Darch, W., Soliman, J., Higgins, N. C., & Morgan, J. T. (2026) · Scientific Reports · doi:10.1038/s41598-026-57258-y · Cited by 1 · see citing works

At the Edge of Survival: Exploring the Frontiers of Tardigrade Extreme Stress Tolerance

Kamilari, M., Pust, F. L., Marcantognini, A., Møbjerg, A., Neves, R. C., Schiøtt, M., & Møbjerg, N. (2026) · Molecular Ecology · doi:10.1111/mec.70471 · Cited by 1 · see citing works

水熊虫耐干燥机制的关键分子筛选

Zheng, K., Shen, P., Hou, Y., Wang, F., Wang, H., & Dong, Y. (2024) · Scientia Sinica Vitae · doi:10.1360/ssv-2023-0270 · Cited by 1 · see citing works

Uncommon N-Glycan Structures in Anhydrobiotic Tardigrades

Yagi, H., Saito, T., Guu, S.-Y., Yamakawa, N., Shimamura, S., Kondo, S., Yagi‐Utsumi, M., Takai, K., Furukawa, J., Guérardel, Y., et al. (2025) · Molecular & Cellular Proteomics · doi:10.1016/j.mcpro.2025.100979 · Cited by 1 · see citing works

Making sense of disorder: Investigating intrinsically disordered proteins in the tardigrade proteome via a computational approachPreprint

Lowe, N. E., & Chang, R. L. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.01.29.478329 · Cited by 1 · see citing works

Room-temperature Storage of Lyophilized Engineered Bacteria using Tardigrade Intrinsically Disordered ProteinsPreprint

Yang, Y., Jiao, Z., Zhang, S., Shan, M., Duan, S., Wang, X., Wang, S., Tang, Y., & Wang, S. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.06.25.449888

Two Novel Heat-Soluble Protein Families Abundantly Expressed in an Anhydrobiotic Tardigrade

Yamaguchi, A., Tanaka, S., Yamaguchi, S., Kuwahara, H., Takamura, C., Imajoh-Ohmi, S., Horikawa, D. D., Toyoda, A., Katayama, T., Arakawa, K., et al. (2012) · PLoS ONE · doi:10.1371/journal.pone.0044209 · Cited by 176 · see citing works

Structural insights into a secretory abundant heat‐soluble protein from an anhydrobiotic tardigrade,Ramazzottius varieornatus

Fukuda, Y., Miura, Y., Mizohata, E., & Tsuyoshi Inoue, A. (2017) · FEBS Letters · doi:10.1002/1873-3468.12752 · Cited by 32 · see citing works

Possible roles of CAHS proteins from Tardigrade in osmotic stress tolerance in mammalian cells

Bino, T., Goto, Y., Maryu, G., Arakawa, K., & Aoki, K. (2024) · Cell Structure and Function · doi:10.1247/csf.24035 · Cited by 4 · see citing works

Tardigrades’ cytoplasmic abundant heat soluble proteins serve as membrane protectors during dehydrationPreprint

Zhang, C. K. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.04.14.648834

Stress-dependent cell stiffening by tardigrade tolerance proteins that reversibly form a filamentous network and gel

Tanaka, A., Nakano, T., Watanabe, K., Masuda, K., Honda, G., Kamata, S., Yasui, R., Kozuka‐Hata, H., WATANABE, C., Chinen, T., et al. (2022) · PLoS Biology · doi:10.1371/journal.pbio.3001780 · Cited by 68 · see citing works