Linking micromorphism,brooding, and hermaphroditism in brachiopods: Insights from Caribbean Argyrotheca (Brachiopoda)

In extant brachiopods, parental brooding of the larvae occurs exclusively within Rhynchonelliformea. Methods of larval protection range from simple retention of the larvae within the mantle cavity, to sophisticated brood care within highly specialized brood pouches found in Argyrotheca and Joania (Terebratulida, Megathyridoidea), Gwynia (Terebratulida, Gwynioidea), and all Thecideoidea (Thecideida). Previous studies on the reproductive biology of Argyrotheca yielded contrasting results on the epithelial origin of the brood pouches in this genus. Here, representatives of different species of Argyrotheca from the Belize Barrier Reef were examined using histological section series. Brood pouches of four species, A. cf. schrammi and Argyrotheca sp. 1–3, are of the same basic structure, formed by invaginations of the anterior body wall and connected to the visceral cavity via the metanephridia. The same four species are simultaneously hermaphroditic, suggesting that fertilization is achieved, at least partly, through selfing. One species, Argyrotheca rubrocostata, differs significantly from all others as it has no brood pouch and gonochoric gonads. Thus, the presence of brood pouches and simultaneous hermaphroditism are concluded to be correlated within Megathyridoidea and proposed to be homologous traits of Joania and several but not all species of Argyrotheca, questioning the monophyletic status of both genera. In contrast to the brood pouches of Thecideoidea, lophophoral epithelium is not involved in the formation of the pouches of Argyrotheca and Joania. Therefore, megathyridoid and thecideoid brood pouches are not homologous but evolved independently within rhynchonelliform brachiopods. All brachiopods with brood pouches share a micromorphic form and a short life span, limiting the space and time available for gamete and larval development. We suggest that the brood pouches and the hermaphroditic gonads of Argyrotheca spp. and Joania compensate these limitations by minimizing the loss of gametes and larvae, and by maximizing the chances of successful fertilization. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Intrinsic vs. extrinsic influences on life history expression: metabolism and parentally induced temperature influences on embryo development rate

Intrinsic processes are assumed to underlie life history expression and trade‐offs, but extrinsic inputs are theorised to shift trait expression and mask trade‐offs within species. Here, we explore application of this theory across species. We do this based on parentally induced embryo temperature as an extrinsic input, and mass‐specific embryo metabolism as an intrinsic process, underlying embryonic development rate. We found that embryonic metabolism followed intrinsic allometry rules among 49 songbird species from temperate and tropical sites. Extrinsic inputs via parentally induced temperatures explained the majority of variation in development rates and masked a relationship with metabolism; metabolism explained a minor proportion of the variation in development rates among species, and only after accounting for temperature effects. We discuss evidence that temperature further obscures the expected interspecific trade‐off between development rate and offspring quality. These results demonstrate the importance of considering extrinsic inputs to trait expression and trade‐offs across species.     

Stock and parental effects on embryonic and early larval development of winter flounder Pseudopleuronectes americanus (Walbaum)

A hierarchical breeding design was used to determine if winter flounder Pseudopleuronectes americanus embryos and yolk-sac larvae sired by Georges Bank males developed and grew larger than fish sired by Passamaquoddy Bay males, and to examine parental contributions to variations in fertilization success, time to 50% hatch, hatch success and larval morphological development. Significant stock effects were detected for time to hatch and larval development. Eggs fertilized by Passamaquoddy Bay males reached 50% hatch significantly earlier than eggs fertilized by Georges Bank males. Larvae sired by Georges Bank males were significantly larger during larval development for four of the six traits measured at 12 days post-hatch: head depth, jaw length, myotome height and body area. Embryo and larval development were strongly influenced by maternal contributions; there were significant maternal variance components for the majority of the variables measured. Paternal variance components were significant for fertilization success, time to hatch, larval jaw length and larval head depth, however, they acted principally through parental interactions. This information has important implications for the long-term sustainable development of winter flounder for aquaculture purposes as well as for understanding winter flounder genetic variation in the wild.     

Regulation of brood size in a burying beetle,Nicrophorus tomentosus (Silphidae)

Regulation of brood size in a biparental burying beetle, Nicrophorus tomentosusWeber, was studied by providing pairs with one of two sizes of mouse carcasses in the laboratory. For a given carcass size, there was an inverse relationship between number and mass offspring in a brood. The requirement for regulation was that brood size was adjusted such that mean mass of individual larvae was constant for carcasses of different size. Brood size was regulated if parents were present but regulation did not occur if parents were removed prior to hatching of larvae. Pairs bred in quick succession on two carcasses raised fewer than the regulated number of young in the second reproductive attempt. Reasons for regulation of brood size in this genus are discussed.     

Monogamy to communal breeding: exploitation of a broad resource base by burying beetles (Nicrophorus)

Abstract.

• 1 To investigate the range of resource size that burying beetles (Nicrophorus) exploit, small (21–33 g), medium (50–90 g) and large (120–210g) carcasses, were placed in the field and then exhumed after 1, 4, 8, 12 or 18 days.
• 2 Nicrophorus attempts to utilize carcasses over this entire size range but has greater success on smaller carcasses.
• 3 Larger carcasses were more difficult to exploit because: (a) they took longer to conceal beneath the leaf litter; (b) they were less likely to be rounded into brood balls; (c) they were more likely to be utilized by dipterans; and (d) they were occupied by greater numbers of congeners.
• 4 Larger carcasses, however, did support greater numbers of larvae and contained broods of greater total mass than smaller carcasses.
• 5 Beetles sometimes bred communally on larger carcasses in the field and same-sex adults were observed to feed young.
• 6 Two follow-up experiments were conducted in the laboratory. On a large carcass N.defodiens, N.tomentosus or N.orbicollis can raise a maximum of 35–50 young. Nicrophorus pustulatus, in contrast, appears to be unique among Nicrophorus in that it can raise nearly 200 young on larger carcasses.
• 7 Nicrophorus orbicollis and Nicrophorus sayi are extremely dependent on parental regurgitations and young fail to survive to the second instar if parents are removed. Young of Nicrophorus defodiens, N.tomentosus and N.pustulatus can develop normally without parental regurgitations.
• 8 I discuss these results in the context of reproduction on carcasses of different size and hypothesize that this breeding system is facultatively quasisocial.

     

Patterns of nest provisioning and parental investment in the solitary digger wasp Ammophila sabulosa

Abstract.

• 1 In nest-building wasps there is a continuum from species that provision each offspring cell with one large prey item (single provisioning) to species that provision each cell with many small items (multiple provisioning). The significance of number of prey per cell was examined during two seasons in Ammophila sabulosa (L.), a species that provisions between one and five caterpillars per cell.
• 2 There was no difference between the total weight of prey in singly-provisioned and multiply-provisioned cells: individual caterpillars placed in multiply-provisioned cells were smaller.
• 3 Small caterpillars were captured and transported to the nest faster than large ones.
• 4 If the first prey provisioned was large, the cell was usually then permanently closed. If the first prey was small, additional prey were captured, and females appeared to become more selective: additional prey were rarely large.
• 5 It is suggested that the flexible provisioning strategy used by A.sabulosa allows a wide range of prey sizes to be utilized without affecting maternal control of offspring size and sex. This could be important when available prey sizes vary temporally and spatially.
• 6 There was a male-biased first generation investment ratio. Female-producing cells were provisioned with a higher total prey weight than male-producing cells, and conversion of prey weight to adult weight was more efficient for females.
• 7 Total prey weight provisioned is probably a good indirect measure of parental time investment in A.sabulosa. Other measures, particularly number of prey provisioned, will be less accurate.

     

Induced reversion of a Chinese hamster ovary triple auxotroph. Validation of the system with several mutagens

A Chinese hamster ovary triple auxotroph (CHO AUXB1) requires glycine, adenosine, and thymidine (GAT) for growth and survival due to a defect in the structural gene for folylpolyglutamate synthetase (FPGS). This auxotroph and others like it contain less than 3% of the parental amounts of FPGS activity. In order to develop a reverse mutation assay with CHO AUXB1, we determined the optimal conditions for measuring reversion and characterized some of the revertants. We also obtained quantitative mutagenicity data for several direct-acting mutagens for comparison to the parental CHO-S/HGPRT locus. Induced revertants appear in the culture immediately following 20-22 h exposures in +GAT complete medium, indicative of dominant genetic changes. They are maximally expressed after 2 population doublings and can be conveniently selected after 44-48 h of expression growth by plating 1 X 10(6) cells/100-mm dish into -GAT-deficient medium and incubating 12-13 days. Plating reconstruction experiments show that the cloning efficiencies of revertants in -GAT medium are not influenced by the presence of up to 1 X 10(6) CHO AUXB1 cells. Dose-dependent increases above the spontaneous revertant frequency (average = 5 X 10(7)) are induced with cis-Pt(NH3)2Cl2 (14-fold) (but not trans-Pt(NH3)2Cl2), PtCl4(10-fold), Pt(SO4)2 (14-fold), K2CrO4 (8-fold), EMS (10-fold), 4-NQO (53-fold), ICR-191 (60-fold), and ICR-170 (30-fold). All of the revertants that have been isolated are stable to repeated subculturing in -GAT medium; 40 out of 42 that have been analyzed are characterized by an increased 72-h growth incorporation of labeled folate and their extracts contain 5-94% as much FPGS as the original, parental CHO-S line. Spontaneous and induced reversion to the GAT+ phenotype primarily reflects mutations involving the FPGS gene locus. But the re-acquisition by most of the revertants of much less than normal amounts of FPGS activity suggests that they arise from compensatory second-site mutations within this gene. Comparison of the mutagenicity patterns of the foregoing compounds as a function of the applied concentration and the relative percent survival reveals some interesting similarities, as well as differences, between the CHO AUXB1/FPGS and CHO-S/HGPRT loci. In particular, the FPGS locus is rather insensitive to EMS (or other simple alkylating agents). However, it seems to be quite susceptible to reversion by other chemicals that are known to react selectively with guanine bases in DNA. CHO AUXBI is a useful supplemental mammalian assay system for assessing quantitatively the generally weak mutagenic activities of metal compounds.     

Functions and social consequences of infant–adult male interaction in a captive group of lowland gorillas (Gorilla gorilla gorilla)

The functions and social consequences of infant-adult male interaction in a captive group of lowland gorillas were evaluated. The two males repeatedly attempted to interact with the infant, the infant sometimes attempted to interact with the males, and the mother usually interrupted interactions between male and infant. Infant-directed actions by the two males frequently showed their interest in the infant and infrequently showed care giving toward her; their other infant-directed actions occurred near the time of excitement or of playful actions between adults. Male-directed actions by the infant frequently showed its interest in one male and infrequently showed care seeking from him. All infant-adult male interactions but one occurred in the less stimulating of the gorillas' two enclosures. Boredom and the mother's frequent thwarting of contact between a male and infant are suggested as influences on the males' interest in the infant. The data suggest that availability of interesting objects alleviated the boredom of captivity for males and thus dissuaded them from seeking stimulation which sometimes resulted in harmful behaviors toward the infant.     

Genetic conflict and evolution of mammalian X-chromosome inactivation

The existence of parentally imprinted gene expression in the somatic tissues of mammals and plants can be explained by a theory of intragenomic genetic conflict, which is a logical extension of classical parent-offspring conflict theory. This theory unites conceptually the phenomena of autosomal imprinting and X-chromosome inactivation. We argue that recent experimental studies of X-chromosome inactivation and andro-genetic development address previously published predictions of the conflict theory, and we discuss possible explanations for the occurrence of random X-inactivation in the somatic tissues of eutherians. © 1995 Wiley-Liss, Inc.     

Parental effort of kestrels (Falco tinnunculus) in nest defense: effects of laying time, brood size, and varying survival prospects of offspring

We studied the nest defense behavior of Eurasian kestrels (Falcotinnunculus) towards a stuffed pine marten (Martes martes) througha 3-year vole cycle (1990–92) in western Finland. Survivalprobability of offspring decreases with a later start of breeding,and, therefore, early breeders should protect their offspringmore than late ones. We found this true for males during theincubation period, but not for females. In addition, we expectedthe nest defense intensity to increase with offspring number.During the incubation period, this was true for females, butnot for males. During the nestling phase, parents did not adjusttheir defense effort to natural or manipulated (by one to twoyoung) brood size. Survival prospects of kestrel offspring werehighest in the increasing vole year 1991 and lowest in the decreasingvole year 1992, and, therefore, we expected the defense activityof kestrels to follow the same trend. However, the oppositeresult appeared true for females with a similar tendency formales. Most hypotheses predicting avian nest defense behaviorwere not supported by our data. Temporally heterogeneous environmentand low degree of nest-site tenacity of migratory kestrels maymake them unfamiliar with environmental variation and survivalprospects of their offspring. Therefore, fitness benefits ofparental care are not predictable, and kestrels may thus adjusttheir parental effort to their own future reproductive potential(i.e., number of future breeding attempts), rather than to somecurrent investment indicator, like offspring age and number.