Background: Stringent regulation of the proteome during periods of stress is essential for the maintenance of cellular homeostasis, function and survival. While proteome regulation has been extensively explored in somatic cells, knowledge of how germ cells respond to proteotoxic stress is lacking.
Methods: This study aimed to investigate how the proteome of mouse spermatocytes and spermatids (n=5 per treatment) are remodelled in response to three proteotoxic stressors; 4-hydroxynonenal (a lipid aldehyde), MG132 (a proteasome inhibitor) and whole-body heat stress (8h at 35°C/16h at 25°C for 14-days). To achieve this, we employed protein solubility fractionation coupled to a high-depth proteomic workflow. This approach permitted the quantification of stress-dependent changes in total protein abundance as well as changes in protein solubility from the same population of cells.
Results: Interrogation of total protein abundance revealed that each proteotoxic stressor induces a unique proteomic signature, with heat stress eliciting the largest response. Further assessment of protein solubility revealed distinct germ cell stage-specific responses to heat stress, with spermatocyte proteins preferentially adopting a ‘less soluble’ state suggesting a shift towards misfolded and/or oligomeric proteoforms while spermatid proteins adopted a ‘more insoluble’ state suggesting accumulation of aggregated proteins. As such, protein aggregation in the germline appears to be an adaptive mechanism, driving the activation of key pathways associated with regulation of the cell cycle, metabolism, transcription and protein homeostasis. Among those proteins identified as potentially aggregating in response to heat stress were several with established roles in spermatogenesis (e.g., heat shock factor protein 5 (HSF5)) and sperm structure/function (e.g., testis-expressed protein 38 (TEX38) and izumo sperm-egg fusion protein 3 (IZUMO3)).
Conclusion: Ultimately, this study provides an increased understanding of stress response pathways in the male germline; a crucial step to inform future strategies to fortify germ cells against stress with implications for fertility and offspring health.