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WIREs Dev Biol
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Peromyscus (deer mice) as developmental models

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Deer mice (Peromyscus) are the most common native North American mammals, and exhibit great natural genetic variation. Wild‐derived stocks from a number of populations are available from the Peromyscus Genetic Stock Center (PGSC). The PGSC also houses a number of natural variants and mutants (many of which appear to differ from Mus). These include metabolic, coat‐color/pattern, neurological, and other morphological variants/mutants. Nearly all these mutants are on a common genetic background, the Peromyscus maniculatus BW stock. Peromyscus are also superior behavior models in areas such as repetitive behavior and pair‐bonding effects, as multiple species are monogamous. While Peromyscus development generally resembles that of Mus and Rattus, prenatal stages have not been as thoroughly studied, and there appear to be intriguing differences (e.g., longer time spent at the two‐cell stage). Development is greatly perturbed in crosses between P. maniculatus (BW) and Peromyscus polionotus (PO). BW females crossed to PO males produce growth‐restricted, but otherwise healthy, fertile offspring which allows for genetic analyses of the many traits that differ between these two species. PO females crossed to BW males produce overgrown but severely dysmorphic conceptuses that rarely survive to late gestation. There are likely many more uses for these animals as developmental models than we have described here. Peromyscus models can now be more fully exploited due to the emerging genetic (full linkage map), genomic (genomes of four stocks have been sequenced) and reproductive resources. This article is categorized under: Establishment of Spatial and Temporal Patterns > Regulation of Size, Proportion, and Timing Nervous System Development > Vertebrates: General Principles Comparative Development and Evolution > Model Systems

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(a) Relationship of Peromyscus relative to other common rodent models. Numbers at nodes indicate estimated divergence dates in millions of years. (b) P. maniculatus (photo: Frances Lee).
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Comparisons of unpublished PO versus BW stock male open field test behavior; data is for 12 PO and 15 BW animals. Animals were place in larger (rat) cage and video‐taped for 5 min. Behaviors (as noted on graph) were recorded at 5 second intervals with the use of the Noldus Observer software (http://www.noldus.com/). The figure shows percentages of time spent in each behavior, minus time spent walking the sides of the cage (which was equivalent between the two species). Bars denote standard error.
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(a) Subadult males resulting from a ♀PO × ♂F1 backcross (♀PO × ♂BW hybrids do not survive) and a ♀bw × ♂po cross, as indicated. (b) and (c) Embryonic fibroblasts from both hybrid crosses (as indicated) at first passage.
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Parental (a) versus ♀PO × ♂BW (b) embryos, roughly equivalent to Mus embryonic day 10 (∼50% are dead by this point). Rare late gestation ♀PO × ♂BW embryos exhibiting hemorrhaging (c), and disproportionately enlarged liver (compare (d) to (e)).
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Selected P. polionotus/P. maniculatus hybrid phenotypes. (a)–(c) are H&E stained sections of mature placentas: (a) PO, (b) ♀bw × ♂po, (c) ♀PO × ♂BW. (d) A ♀PO × ♂BW molar conceptus (i.e., apparently all extra‐embryonic tissues).
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Additional Peromyscus mutants/variants: (a) Tan streak (tns); (b) Ashy (ahy) mutants of varying ages (5–12 months) exhibiting varying degrees of the phenotype; (c) P. polionotus subgriseus (PO stock; right) compared to P. polionotus leucocephalus (LS stock; left). (d) Cataract‐webbed (cwb) exhibiting cataracts; and (e) cwb—foot with webbed toes. Photos: Frances Lee, except (c) Clint Cook.
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Origins of PGSC stocks; species and two‐letter stock designation indicated.
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Selected Peromyscus mutants/variants with visible external phenotypes: (a) Dominant Spotting (S). (b) Variable White (Vw). (c) Nonvariant P. maniculatus bairdii (BW stock). (d) Wide Band Agouti (ANb). (e) Platinum (plt). (f) Blonde (bln). (g) albino (c; left) and ivory (i; right). (h) Hairless 2 (hr‐2; left—reduced vibrissae) and Hairless 1 (hr‐1; right—WT vibrissae). Photos: Frances Lee.
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Selected developmental stages of Peromyscus maniculatus BW stock. (a) Flushed two‐cell embryos. (b) Flushed embryos 48 h after artificial insemination, cultured in KSOM. (c) Blastocysts. (d) Embryonic day 14 Embryo (∼ to Mus E10). (e) Embryonic day 17 Embryo (∼ to Mus E13). (f) Embryonic day 20 Embryo (∼ to Mus E16).
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Nervous System Development > Vertebrates: General Principles
Establishment of Spatial and Temporal Patterns > Regulation of Size, Proportion, and Timing
Comparative Development and Evolution > Model Systems