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Maternally Inherited Peptides As Strain-Specific Chemosignals

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Maternally Inherited Peptides As Strain-Specific Chemosignals Maternally inherited peptides as strain-specific chemosignals Hideto Kabaa,1,2, Hiroko Fujitaa,1, Takeshi Agatsumab, and Hiroaki Matsunamic,d,e,2 aDepartment of Physiology, Kochi Medical School, Kochi University, Nankoku, 783-8505 Kochi, Japan; bDepartment of Environmental Health Sciences, Kochi Medical School, Kochi University, Nankoku, 783-8505 Kochi, Japan; cDepartment of Molecular Genetics and Microbiology, Duke University Medical Center, Durham, NC 27710; dDepartment of Neurobiology, Duke University Medical Center, Durham, NC 27710; and eDuke Institute for Brain Sciences, Duke University, Durham, NC 27710 Edited by Frank Zufall, Saarland University, Homburg, Germany, and accepted by Editorial Board Member John R. Carlson October 13, 2020 (received for review July 13, 2020) Most mammals rely on chemosensory cues for individual recogni- of the ESP1 receptor, V2Rp5, which is expressed in vomeronasal tion, which is essential to many aspects of social behavior, such as sensory neurons (VSNs). maternal bonding, mate recognition, and inbreeding avoidance. We have focused on mitochondrially encoded peptides as a Both volatile molecules and nonvolatile peptides secreted by candidate source of individual chemosignals, based on highly individual conspecifics are detected by olfactory sensory neurons polymorphic genes. Because NADH dehydrogenase 1 (ND1) in the olfactory epithelium and the vomeronasal organ. The perti- and NADH dehydrogenase 2 (ND2) are encoded in the mito- nent cues used for individual recognition remain largely unidenti- chondrial genome, they are maternally inherited, formylated at fied. Here we show that nonformylated, but not N-formylated, the N-terminal methionine at the time of synthesis, and poly- mitochondrially encoded peptides—that is, the nine N-terminal morphic among inbred strains (Fig. 1). The ND1 peptide is de- amino acids of NADH dehydrogenases 1 and 2—canbeusedto rived from the nine N-terminal amino acids of the ND1 protein: convey strain-specific information among individual mice. We dem- MFFINXLTL. The identity of the sixth amino acid varies among onstrate that these nonformylated peptides are sufficient to induce different strains and can be either A, I, T, or V. ND2 has similar a strain-selective pregnancy block. We also observed that the preg- amino acid substitutions. These features raise the possibility that nancy block by an unfamiliar peptide derived from a male of a dif- mitochondrially encoded peptides can be used to convey mouse ferent strain was prevented by a memory formed at the time of strain-specific information among individuals. NEUROSCIENCE mating with that male. Our findings also demonstrate that This possibility can be explored in the context of the selective pregnancy-blocking chemosignals in the urine are maternally pregnancy block effect, which is also known as the Bruce effect inherited, as evidenced by the production of reciprocal sons from (21) and depends on a functional vomeronasal system (22–25). two inbred strains and our test of their urine’s ability to block preg- In this behavioral paradigm, a high incidence of pregnancy fail- nancy. We propose that this link between polymorphic mitochon- ure occurs when recently mated female mice are exposed to drial peptides and individual recognition provides the molecular means to communicate an individual’s maternal lineage and strain. chemosignals from a second unfamiliar male. The female re- ceives the signal from an unfamiliar male through her VNO, mitochondria | olfaction | maternal inheritance | pheromone | Bruce effect resulting in a suppression of the prolactin and progesterone Significance he ability to distinguish and recognize individuals is critical Tfor animals’ adaptive regulation of social behavior. Although individual information may be coded in a number of ways, che- For most mammals, olfaction plays an important role in indi- mosensory cues are a primary means by which many animals vidual recognition. The social chemosignals secreted by indi- evaluate others’ identity, relatedness, sex, and social status (1–6). vidual conspecifics are complex variable mixtures that are still Each individual animal has a great complex of chemosensory poorly understood. Here we identify nonformylated, but not N-formylated, mitochondrially encoded peptides: That is, the cues, some self-generated and others acquired from the envi- nine N-terminal amino acids of NADH dehydrogenases 1 and 2 ronment and conspecifics (7). The resulting chemical signatures as reliable chemosignals for individual recognition by using a are complex variable mixtures that are still poorly understood. vomeronasal organ-mediated pregnancy block, known as the Three highly polymorphic multigene families in mice are Bruce effect. Our findings also revealed that pregnancy- known to contribute to individual differences in chemosensory blocking chemosignals in the urine are maternally inherited. cues that are detected through specific vomeronasal receptors We propose that this individuality cue provides the molecular expressed in sensory neurons of the vomeronasal organ (VNO). means to communicate an individual’s maternal lineage The major histocompatibility complex (MHC) genotype has and strain. been linked to mate recognition in the pregnancy block effect, as congenic mice differing from the mating male only in their MHC Author contributions: H.K. and H.M. designed research; H.K. and H.F. performed research; genotype were effective in blocking pregnancy (8). Further evi- H.K., H.F., T.A., and H.M. contributed new reagents/analytic tools; H.K. and H.F. analyzed dence has shown that the MHC peptide ligands themselves data; and H.K., H.F., T.A., and H.M. wrote the paper. Competing interest statement: H.M. receives royalties from Chemcom. Other authors convey strain-specific information in the context of the preg- declare no competing interest. nancy block effect (9, 10). Major urinary proteins provide an This article is a PNAS Direct Submission. F.Z. is a guest editor invited by the individual genetic identity signature that underlies individual Editorial Board. – recognition, territorial behavior, and mate choice (11 16). Published under the PNAS license. A series of studies demonstrated that exocrine-gland secreting 1H.K. and H.F. contributed equally to this work. peptide 1 (ESP1) that is released into the tear fluids of adult 2 To whom correspondence may be addressed. Email: [email protected] or hiroaki. male mice is one of the key factors that cause pregnancy block [email protected]. – (17 20). Female mice exhibit high pregnancy failure rates upon This article contains supporting information online at https://www.pnas.org/lookup/suppl/ encountering males that secrete different levels of ESP1 com- doi:10.1073/pnas.2014712117/-/DCSupplemental. pared to the mated male. This effect is mediated via the activation www.pnas.org/cgi/doi/10.1073/pnas.2014712117 PNAS Latest Articles | 1of6 Downloaded by guest on October 2, 2021 male urine at a final concentration of 50 μM and testing its pregnancy-blocking effectiveness following mating with a BALB/ c male. The N-formylated variants of the ND1 and ND2 peptides and the prototypical bacterial N-formyl-methionyl-leucyl-phe- nylalanine (MLF) were ineffective at inducing pregnancy failure (Fig. 3A). N-formylated ND1-6A was still ineffective even when applied at a high dose (150 μM). The low rate of pregnancy failure in the group exposed to the N-formylated ND1 or ND2 variant is not ascribed to an overall decrease in the susceptibility to pregnancy block, because pregnancy-blocking effectiveness Fig. 1. N-terminal polymorphisms in the mouse mitochondrial ND1 and was confirmed by exposure to CBA male urine that served as a ND2 proteins. The polymorphic positions in ND1 and ND2 are shown in bold. positive control. Parentheses indicate H-2 haplotypes, where known. f-M, formyl-methionine. In contrast, the nonformylated forms of ND1-6A and ND2-7T derived from the NZB strain significantly increased the effec- tiveness of BALB/c male urine in blocking pregnancy, whereas levels that are necessary for embryonic implantation. Hence, the the nonformylated form of ND1-6I or ND2-7A derived from the presence of an unfamiliar male can terminate pregnancies by a BALB/c strain did not (Fig. 3B). In addition, the pregnancy block male of a different strain, but it does not block pregnancies that effect of nonformylated ND1-6A was dose-dependent; a low he himself (or a mouse of the identical strain) has initiated. This μ B “ ” is because the female learns to recognize and remember her dose (5 M) was without effect (Fig. 3 ). Thus, familiar urine “ ” mate’s chemosignals during a sensitive period at the time of could be converted to unfamiliar urine by the addition of a mating and gates their effects in order to maintain the pregnancy. single nonformylated peptide derived from a strain other than BALB/c. The potency of nonformylated ND1-6A in blocking Results pregnancy was comparable to that of AAPDNRETF (9), an b N-Terminal Polymorphisms in the ND1 and ND2 Proteins. The mito- MHC class I peptide of the disparate H-2 haplotype (Fig. 3B). A chondrial DNA sequencing confirmed that the inbred strains of scrambled version of nonformylated ND1-6A MFFINALTL, mice (i.e., BALB/cCrSlc, CBA/NSlc, and NZB/NSlc) from Japan namely LMATLFNIF, failed to induce pregnancy failure, suggesting SLC have the same N-terminal amino acid sequences deduced from the nucleotide sequences of the mitochondrial nd1 and nd2 genes as the corresponding inbred strains
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