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During spermatogenesis, mitochondria extend along the whole length of spermatid tail and offer a structural platform for microtubule reorganization and synchronized spermatid individualization, that eventually helps to generate mature sperm in Drosophila. However, the regulatory mechanism of spermatid mitochondria during elongation remains largely unknown. Herein, we demonstrated that NADH dehydrogenase (ubiquinone) 42 kDa subunit (ND-42) was essential for male fertility and spermatid elongation in Drosophila. Moreover, ND-42 depletion led to mitochondrial disorders in Drosophila testes. Based on single-cell RNA-sequencing (scRNA-seq), we identified 15 distinct cell clusters, including several unanticipated transitional subpopulations or differentiative stages for testicular germ cell complexity in Drosophila testes. Enrichments of the transcriptional regulatory network in the late-stage cell populations revealed key roles of ND-42 in mitochondria and its related biological processes during spermatid elongation. Notably, we demonstrated that ND-42 depletion led to maintenance defects of the major mitochondrial derivative and the minor mitochondrial derivative by affecting mitochondrial membrane potential and mitochondrial-encoded genes. Our study proposes a novel regulatory mechanism of ND-42 for spermatid mitochondrial derivative maintenance, contributing to a better understanding of spermatid elongation., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

Lonchoptera lutea males produce giant spermatozoa that are more than 2000 μm long and 1.4 μm wide. Unlike the typical brachyceran spermatozoon, they have a highly asymmetrical cross-section with only a single, albeit very large, mitochondrial derivative and a pair of massive accessory bodies, one of which extends throughout the entire length of the sperm tail. The accessory bodies consist of an electron-dense matrix in which numerous peculiar electron-lucid substructures are embedded. In the mated female, the giant spermatozoa are found inside two tubular spermathecae which are also extremely long, measuring 4000 μm or more., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

The spermatozoa of Diasemopsis comoroensis and Diasemopsis meigenii differ from the conventional brachyceran type in several respects. Not only are they very long but they are also extraordinarily wide, especially at the very end of the tail. The latter is effected by two cellular components: oversized mitochondrial derivatives and a prominent central band, which is a peculiar structure not known from any other spermatozoa. Based on its position with respect to the other organelles and its origin during early spermiogenesis, the central band is interpreted as a derivative of the centriolar adjunct material. Like the axoneme, the mitochondrial derivatives and the central band extend through the entire length of the spermatozoon tail. The spermatozoon tail is helical and can be coiled up into a tight cone-shaped spiral with a peculiar corkscrew shaped end portion. The potential adaptive significance of these features and their coevolution with the morphology of the female's multi-chambered ventral receptacle are discussed, as are sperm competition and cryptic female choice., (Copyright © 2015 Elsevier Ltd. All rights reserved.)