Milk composition during lactation suggests a mechanism for male biased allocation of maternal resources in the tammar wallaby (Macropus eugenii)

Abstract

Recent research has found empirical evidence in support of the Trivers-Willard Hypothesis that offspring sex allocation is correlated with maternal investment. Tammar wallabies birthing sons have higher investment ability; however a mechanism for sex specific differential allocation of maternal resources in wallabies remains elusive. In metatherians the majority of maternal investment is during lactation. To examine if differential allocation occurs during lactation, we measured total milk protein, lipid and carbohydrates, from mothers with male and female pouch young, during phase 2B (100-215 days post partum) and phase 3 (215-360 days post partum) of lactation. Mothers of sons allocated significantly higher levels of protein than mothers of daughters during phase 2B of lactation, however no sex specific difference in maternal allocation was found for lipids, carbohydrates, or any milk component during phase 3 of lactation. We were unable to measure milk production to establish any differences in the amount of milk allocated. However, with the production of more milk comes a dilution effect on milk components. Given that we find no apparent dilution of milk components may suggest equality in milk production. Offspring body weight at 14 months of age was related to protein allocation during phase 2B of lactation, providing a maternal mechanism for differential allocation with fitness consequences. We believe collection of earlier phase 2A (0-100 days post partum) milk may yield important results given that differential investment in metatherians may be most apparent early in lactation, prior to any significant maternal investment, when a decision on termination of investment can be made with very little energetic loss to the mother. Interestingly, small mothers did not birth sons and better maternal condition was associated with raising sons. These data are in support of TWH and demonstrate a potential mechanism through which condition dependent and sex specific maternal investment may occur.

Figure 1. Milk components by offspring sex.

(a) protein (b) carbohydrate and (c) lipid content in milk during phase 2B and phase 3 of lactation in wallabies birthing sons (shaded) or daughters (unshaded). Females allocate significantly more protein to sons during Phase 2B of lactation than to daughters, while females allocate both carbohydrates and lipids equally to sons and daughters during phase 2B and 3 of lactation. Levels not connected by the same letter are significantly different (post-hoc Student’s t-test).

Figure 2. Gross milk energy content during phase 2B and phase 3 of lactation.

Mean changes (± s.e.) in the energy content of milk from Tammar wallaby mothers raising sons (shaded bars) and those raising daughters (open bars) during phase 2B and phase 3 of lactation. Despite the allocation of different milk components the total energy content of the milk is not significantly different between the milk allocated to sons and daughters.

Figure 3. Smaller females only produce daughters.

Maternal body size (pes length) and offspring sex. Females with a pes length of less than 139 mm only produced daughters. Although, mothers pes length did not predict offspring sex (Logistic regression: ChiSq = 2.08, p = 0.151).

Figure 4. Protein allocation during phase 2B of lactation influences offspring weight into adulthood.

Maternally allocated proteins during phase 2B of lactation influences offspring body weight at fourteen months of age.