Department of Genetics

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Assessment of yield traits between family groups of the cultured abalone (Haliotis midae) in South Africa
(Stellenbosch : Stellenbosch University, 2012-03) Van Schalkwyk, Hester Josina; Brink, Danie; Stellenbosch University. Faculty of AgriSciences. Dept. of Genetics.
ENGLISH ABSTRACT: The abalone Haliotis midae is the most important aquaculture species in South Africa. The industry is dependent on export to Far Eastern markets in a variety of forms, including live, frozen, canned or dried. The species is considered undomesticated in the sense that the current commercial broodstock has been obtained from natural populations through a process of random collection. Global competition has necessitated the South African industry to introduce a genetic improvement program to increase biological productivity and financial profitability. The objective of this study was to assess the genetic variation and to estimate key parameters in terms of growth and yield related traits, between family groups that form part of the breeding program. The study reports on heritability estimates of growth rate (0.14 ± 0.05), canning yield (0.08 ± 0.03), and drip loss during live export (0.03 ± 0.02). The high genetic correlation (0.94 ± 0.34) between shell length and live weight enables industry to utilise either weight or shell length as a criteria during operational practices such as sorting, grading and harvesting. The correlation of 0.85 ± 0.01 between live weight and canning loss indicates that animals that weigh more have a lower dressing percentage. Based on these low heritability values obtained for yield related traits it is recommended not to include these traits in the selection program at this stage. The findings of the study were however, compromised by the availability of a limited number of family groups, the age differences between families and the effect of different locations on the variance in phenotypes. Further investigation is needed to confirm the credibility of the results.
A comparative analysis of growth traits in Triploid and Diploid Genotypes of the South African abalone, Haliotis midae
(Stellenbosch : Stellenbosch University, 2011-12) Prins, Nico; Brink, Danie; Stellenbosch University. Faculty of AgriSciences. Dept. of Genetics.
ENGLISH ABSTRACT: Abalone production is the largest financial contributor to aquaculture in South Africa and practically all of the abalone produced is exported to Asia. This means that the product must be globally competitive and many technologies have been applied to this cause. One that specifically shows great promise for bivalve mollusc production is triploidy; more precisely, sterility due to the induction of aneuploidy. Under normal maturation, energy is diverted from somatic growth through sexual maturation, therefore inhibiting or retarding gametogenesis through a process such as aneuploidy is expected to increase growth and decrease the time to marketing. Two studies preceding this one investigated the induction of triploidy through hydrostatic shock (De Beer, 2004) and the comparative growth rate of triploid genotypes from 8 to 24 months, prior to the onset of sexual maturation (Schoonbee, 2008). During this comparative growth stage, no convincing statistical evidence of faster growth or of seasonal environmental effects could be obtained. It was recommended that growth between triploid and diploid variants be compared during the age period when sexual maturity becomes a factor to determine whether triploidy in Haliotis midae is a useful biotechnological tool to improve biological productivity and global competiveness of the abalone industry. The growth measured as shell length and wet weight in the period from 29 to 62 months showed a statistically significant difference in mean weight and mean length with diploids showing a superior growth rate compared to their triploid siblings. This difference of 1.99 mm and 5.13 g was however not perceived as being commercially significant. Important production parameters including canning yield percentage and gonadosomatic index were also measured during this trial. For both these parameters, the triploid genotype showed statistically and commercially significant improvement of 10.68% increased canning yield and 28.42% reduction in gonadosomatic index when compared to their diploid counterparts. Triploid abalone was found to be not completely sterile; gametes and even mature gonads were observed in some instances. Even though complete sterility was not achieved there appeared to be a retarded gonadosomatic development in triploid variants. The delay in sexual maturation, together with the improvement in canning yield, may justify triploidy’s commercial application, despite its reduced growth rate.
Critical assessment of the “internal reference” method to eliminate non-genetic effects within a Combined Family Selection program on the abalone species (Haliotis midae)
(Stellenbosch : Stellenbosch University, 2013-12) Difford, Gareth Frank; Brink, Danie; Stellenbosch University. Faculty of Agricultural Science. Dept. of Genetics, division of aquaculture.
ENGLISH ABSTRACT: The aim of this study was to critically assess the implementation of the internal reference method within the most recent 173 full-sibling growth trial of the Innovation Fund Abalone Breeding Project. The trial was conducted over two locations for a period of five years, with minimal replication for the majority of test families and a single full-sibling family was entered into each experimental unit (basket) as an internal reference group. The primary focus was firstly, to validate the performance of the internal reference group as a control for comparisons and correction of environmental variation in test family performances. Secondly, to identify areas of weakness and either make recommendations to remedy areas of weakness or justify devoting resources to alternative methods of reducing extraneous environmental variance with limitations on replication. The efficiency and statistical power associated with utilising internal reference information as a covariate and for manual correction respectively were examined for the 6 full-sibling test families that were replicated. This study reports on the evaluation of factors which are potential sources of bias in the internal reference method, the first of which, tag loss, was found to be significant after 6- 12 months. However, it was not found to bias internal reference group performances as the factors which contribute to tag loss were found to act randomly. Variability in size ratio of internal reference to test family at co-stocking proved a significant source of bias, as reference groups smaller than their test family counterparts had reduced performances. Testing for genotype by environment interactions was precluded due to the inherent lack of replication and the subsequent confounding of genotype effects with inter-rearing structure effects at one of the locations. However, significant differences were detected for both traits of interest of the internal reference group over the two locations. Significant antagonistic interactions were detected and identified as a source of bias for average daily weight gain of replicate test families. The evaluation of average daily length gain for the efficiency of adjustment when the internal reference is a covariate and the change in statistical power when the internal reference is used for a manual correction, yielded conflicting results. The latter shows a decrease in statistical power and the former shows an increase in efficiency, both resulting in poor goodness of fit in the respective models. There was however evidence that when no antagonistic interactions occurred “between replicate variance” decreased and therefore the internal reference method has statistical merit provided all critical success factors are satisfied. Recommendations were made for future implementation of the internal reference method to facilitate adequate statistical testing for sources of bias and the prevention thereof. Additionally, an alternative method which may have merit in decreasing environmental variance and the need for replication, is discussed.
The effect of triploidy on the growth and survival of the indigenous abalone, Haliotis midae, over a 24 month period under commercial rearing conditions
(Stellenbosch : Stellenbosch University, 2008-03) Schoonbee, Lize; Brink, Danie; Stellenbosch University. Faculty of AgriSciences. Dept. of Genetics.
ENGLISH ABSTRACT: Triploidy is the genetic state of containing three sets of chromosomes per cell in stead of two as in diploid organisms. The South African abalone (Haliotis midae) is naturally a diploid organism that sexually matures between four to eight years of age. Early sexual maturity is a disadvantage in cultured abalone stock, as the process of gonad development and spawning is energy demanding, causing energy to be diverted away from somatic growth. This same problem has been extensively experienced in diploid bivalve molluscs, where triploidy has since been applied as a means to prevent sexual maturation from occurring, thereby speeding up the growth process and shortening the time to marketing. Because triploidy was effective in bivalves, it was thought that it could contribute to faster growth in abalone as well. A procedure for the induction of triploidy in the abalone, Haliotis midae, was developed by De Beer (2004) and yielded up to 100 percent triploidy in treated abalone larvae. The next step was to compare the growth of the diploids and triploids to establish whether there was indeed a growth advantage on the part of the triploids, in view of commercial application. By using the same techniques as described by De Beer (2004), three groups consisting of triploid and diploid siblings were produced and subscribed to a comparative growth trial. The groups were spawned in three different seasons. The main objective was to establish whether there was in fact a difference in growth between diploid and triploid siblings, and whether seasonal effects were associated with growth advantages for either triploids or diploids. The two growth parameters measured were shell length and body weight. Measurements commenced at eight months of age, when the abalone could be individually tagged and continued up to the age of 24 months. The over-all results provided no convincing evidence of statistically significant faster growth of triploid juveniles compared to that of diploids up to two years of age. Growth differences were detected between seasons, but could not confidently be ascribed to seasonal environmental effects. The regression of shell length to body weight was similar for diploids and triploids.
Genetic assessment of five breeding populations of abalone (Haliotis midae) through a comparative Performance Testing Scheme
(Stellenbosch : Stellenbosch University, 2015-04) Vlok, Arnoldus Christiaan; Brink, Danie; Stellenbosch University. Faculty of Agrisciences. Dept. of Genetics.
ENGLISH ABSTRACT: Cultured abalone in South Africa is undomesticated. For the local industry to remain competitive on the international markets it is essential to improve production. This study is part of a selective breeding component of a larger genetic programme that aims to enhance productivity of the local industry by genetic improvement of growth rates. Selective breeding programmes are based on genetic variation and correlations. Molecular studies proved genetic differentiation exist between the broodstock- and offspring populations and among the offspring populations used in this study. Five commercial abalone farms from the Walker bay region each entered 3000 randomly selected animals obtained from synchronised mass spawning of conditioned broodstock into a Performance Recording Scheme (PRS). Microsatellite marker analysis proved these broodstock populations to be representative of the wild populations. The five cohorts were assessed over the five locations represented by three replicates per location with 200 randomly assigned animals per replicate. The average growth rate was used as growth performance parameter by measuring shell length and body weight at three month intervals over a period of 24 months. Interaction was observed between cohort and location effects when analysing the full data set. This was unexpected as the cohorts were constructed from parent stick that was randomly sampled from the same geographical area, the larger Walker bay. The factors suspected of causing this observed interaction were considered in a stepwise analysis. Initial and progressive tag loss, differences in initial size of animals entered into the study and on-farm management errors were considered as possible causes of the observed interaction in a stepwise analysis. Statistically significant differences were observed between the five cohorts and between the five locations in terms of length and weight growth rates. Based on these findings it is advised that a central facility is used to effectively compare the growth rates of different cohorts or populations. Any future research in selective breeding to follow this study should involve the integration of molecular techniques and biotechnologies.
Identification of molecular markers for the diagnostic identification of the intracellular prokaryote associated with the appearance of withering syndrome in the abalone Haliotis midae
(Stellenbosch : University of Stellenbosch, 2005-03) Ockert, Candice; D'Amato, M. E.; Brink, Danie; University of Stellenbosch. Faculty of Agrisciences. Dept. of Genetics.
Withering syndrome is a severe disease of abalone, Haliotis species that has been associated with mortality ranging from 99% in black, H. cracherodii Leach and 30% in red abalone, H. rufescens Swainson. The disease was first observed in California, along the west coast of North America and is an economically important disease that has led to the closure of the black abalone fishery throughout the southern California State. The causative agent of withering syndrome is a gram-negative intracellular Rickettsiales-like prokaryote designated Candidatus xenohaliotis californiensis. The geographical range of C. xenohaliotis californiensis is broad, besides red and black abalone it has also been reported in yellow, H. corrugate and blue abalone, H. fulgens all caught in Baja California, Mexico. In 2000 a Rickettsiales-like prokaryote resembling the disease-causing agent was observed in the digestive gland of the South African abalone H. midae. In this study we aimed to determine the relationship of the bacterium observed in the local abalone species, H. midae to the disease-causing agent of withering syndrome. A specific PCR and in situ hybridization test using primers and probes specific for the C. xenohaliotis californiesis 16S rDNA gene were used to screen H. midae digestive gland tissues showing classical features of the Rickettsiales-like prokaryote. Both analyses indicated that C. xenohaliotis californiensis is not present in the local abalone species. We therefore aimed the identification of the organism parasiting the local abalone species by DNA sequence analysis of the 16S rDNA gene. The 16S rDNA gene was amplified by PCR, cloned and sequenced. Phylogenetic trees, constructed by maximum parsimony analysis revealed a diverse community comprised of α - Proteobacteria, Mollicutes and Spirochaetes. In the class α - Proteobacteria a novel group of sequences showing phylogenetic affinities to the order Rickettsiales was identified as likely candidate for forming the Rickettsiales-like inclusions in the digestive gland of H. midae. Oligonucleotide probes that bind to four variable regions of the novel group were used to confirm their presence in infected H. midae digestive gland tissue by in situ hybridization. Although these probes did not recognize the inclusions formed by the Rickettsiales-like organisms, they revealed the presence of a group of free-living bacteria abundant in the host tissue. We therefore conclude that (1) C.xenohaliotis californiensis is not present in the South- African abalone, H. midae; (2) the organisms isolated from the digestive gland of H. midae are part of the normal microflora and (3) the group of sequences showing phylogenetic affinities to the order Rickettsiales is not responsible for the Rickettsiales-like inclusions in infected digestive gland tissues but represent a novel group of organisms that are abundant in the host tissue.
Induction of triploidy in the South African abalone, Haliotis midae, by the use of hydrostatic pressure
(Stellenbosch : University of Stellenbosch, 2004-12) De Beer, Mathilde; Brink, Danie; University of Stellenbosch. Faculty of Agrisciences. Dept. of Genetics.
ENGLISH ABSTRACT: The indigenous abalone, Haliotis midae has been a successfully cultured aquaculture species in South Africa since 1990. It has a slow growth rate and takes from two to five years to reach market size. Like for most other commercially important abalone species, the slow growth rate of H. midae is a cause of concern with regard to the profitability of farming and global competitiveness of the species. Ploidy manipulation of the maternal genome, a universally growing practice in shellfish culture, is considered a promising method to improve the growth rate of abalone - a desirable trait in aquaculture organisms from a commercial perspective. This manipulation technique is employed to achieve sterility, which results in limited gonad development. The consequent re-allocation of resources to somatic growth results in improved growth. The purpose of this study was to establish a viable method for the induction and validation of triploidy, on a commercial scale, in the South African abalone, H. midae. The focus was on hydrostatic pressure as a method of induction and flow cytometry as the method of validation. The results obtained confirm hydrostatic pressure as an effective method for the induction of triploidy in H. midae, delivering high percentages of triploidy (>80%) over a wide range of pressures and times, in 48 hour-old larvae. Hydrostatic pressure had a negative effect on survival in 20 hour-old larvae. Flow cytometry was validated as a reliable, fast and accurate, though expensive, method for identification of triploidy in H. midae. As an outcome of this study a manual of “Procedures for the Induction and Validation of Triploidy in the abalone” is presented (Appendix 1) together with recommendations for further studies on triploidy in the South African abalone, H. midae.
Microsatellite marker development and parentage assignment in Haliotis midae
(Stellenbosch : Stellenbosch University, 2008-03) Van den Berg, Nicol-Candice; Roodt-Wilding, R.; Brink, Danie; Stellenbosch University. Faculty of Agrisciences. Dept. of Genetics.
ENGLISH ABSTRACT: The five leading abalone producers in South Africa have initiated a genetic enhancement program for Haliotis midae in a collaborative effort to improve economically valuable traits. Several independent objective-specific studies were initiated, including the establishment of a Performance Recording Scheme (PRS), utilised in this study, and necessary to monitor the ongoing performance of individuals as the move from mass-selection to marker assisted selection (MAS) is implemented. The primary objective of this study was parentage assignment of F1 offspring mass-selected for size at approximately one year and allocated to either a “faster” or a “slower” growth group. Nine microsatellite markers were used to genotype juveniles and potential parents, with assignment completed using CERVUS 2.0. Average growth results for Abagold and HIK were comparable for both growth groups. Slight environmental effects, although not statistically significant, were evident as growth advantages for juveniles within the faster growth group at two of the five locations and for juveniles within the slower growth group at one of the five rearing locations. Despite measures to standardise environmental influences, variables are difficult to control within the reality of a production environment; and potential genotype x environment interactions may require further investigation and factoring into future breeding programs. The additional costs associated with MAS often make the technology prohibitive to most aquaculture operations, despite the significant genetic gains to be realised from its implementation. Cost-optimising routine processes such as DNA extractions may be one approach to reduce these additional costs. Chelex®100 appears to be a suitable alternative to the CTAB method – being quick and cost-effective to perform. Applying this method in combination with the high throughput of a robotic platform warrants further evaluation. For the microsatellite development, 50% of positive recombinant clones contained inserts. Sequencing of these clones produced 16% perfect repeats and 47% imperfect repeats for which 52 primer sets were designed and tested. In total, 31 polymorphic microsatellite loci of different motifs and composition were developed. Sixty-one percent of sequenced clones were deemed redundant and pre-screening for both uniqueness and the presence of microsatellites would reduce unnecessary sequencing thus improving the efficiency of the FIASCO method and reducing costs. Nine loci were selected for parentage assignments. Null alleles were present for all the selected markers; however, frequencies were below the critical level of 5%. Parentage yielded 91% and 90% successful assignment for Abagold and HIK respectively; however, observations indicate that a measure of relatedness may exist between breeders. Recommendations with regards to future family breeding include, for both Abagold and HIK, retaining selected breeders based on their respective contributions to the F1 progeny while reassessing the potential of remaining breeding stock under more controlled breeding conditions. No obvious trends were observed for growth with most individuals producing both faster and slower growing offspring. Juveniles will be reassessed at two years to determine whether the size advantage or disadvantages were maintained and to ascertain whether growth advantages/disadvantages may be gender specific.
Molecular analysis of genetic variation and relationships amongst abalone (Haliotis midae) at the I&J abalone hatchery at Danger Point, Gansbaai, R.S.A.
(Stellenbosch : Stellenbosch University, 2002-12) Lambrechts, Daneel; Brink, Danie; D'Amato, M. E.; Stellenbosch University. Faculty of AgriSciences. Dept. of Genetics.
ENGLISH ABSTRACT: The species Haliotis midae is the only abalone species of commercial importance to the South African fisheries industry. The species is currently under substantial pressure due to commercial harvest and illegal poaching, to the extent that genetic and biodiversity may be under threat. The species is also being cultured in commercial systems for supply to international markets. The South African production for 2002 amounted to 350 tons. The commercial production of abalone may also have implications with regard to genetic diversity of natural and commercial populations. The aim of this project was to assess the genetic diversity of the natural and commercial populations of abalone at the I&J Abalone Farm, Danger Point, Gansbaai, in South Africa, in order to make recommendation with regard to possible impact on natural diversity as well as internal management systems. Distortion of natural genetic diversity or a loss of internal diversity will have detrimental consequences. Representative samples have been obtained from the surrounding natural population, as well as the commercial populations, including the broodstock and various progeny groups. Both mtDNA and AFLP molecular techniques were used to assess genetic diversity. Data analysis showed that the genetic profile of the commercial population display sufficient genetic variation. The genetic structure of the commercial population also displays no significant deviation from that of the surrounding natural population, i.e. the population of origin. The results through give indication of a small, though unsignificant loss of genetic variation from the broodstock to the subsequent progeny groups. The investigation conclude that the commercial populations of abalone at the I&J Abalone Farm, Danger Point, Gansbaai, in its current format holds no threat to the disruption of the genetic diversity of the surrounding natural population. Further, the commercial population possesses sufficient genetic variation in view of future genetic development. There is signs of a slight loss of genetic variation in the change over from the broodstock to the progeny groups. This would necessitate careful and controlled replacement of the original broodstock with new animals from the commercial progeny groups as part of an ongoing process of domestication.
Quantitative assessment of yield traits between family groups of the cultured abalone, Haliotis midae, during the process of canning
(Stellenbosch : Stellenbosch University, 2013-03) Gerber, Maria Elizabeth (Mariette); Brink, Danie; Stellenbosch University. Faculty of AgriSciences. Dept. of Genetics.
ENGLISH ABSTRACT: The species Haliotis midae is of great commercial value to the South African abalone industry and is mainly exported to Asian markets, specifically China. Up to 50% is sold as canned products with H. midae registering an average canning yield of approximately 35%. The species is presently genetically undomesticated and breeding programmes are being introduced to improve a range of production traits of which growth and yield is of primary importance. The objective of the study was to determine genetic parameters such as heritability, genotypic and phenotypic correlations of yield-related traits to assess the potential genetic improvement through selective breeding. A series of yield-related parameters were identified that is of relevance to the standard abalone canning procedure. Low to moderate heritabilities where recorded for most traits, including pre-shuck/live weight (0.20 ± 0.06), post-shuck weight (0.15 ± 0.05), post-gut weight (0.15 ± 0.05), post-brine weight (0.19 ± 0.06), pre-canning weight (0.19 ± 0.06), post-canning weight (0.21 ± 0.06), shell weight (0.16 ± 0.05), canning yield percentage (0.08 ± 0.03) and shell weight to post-gut weight ratio (SW: PGW) (0.09 ± 0.04). Weight related parameters are phenotypically highly correlated (0.86 ≤ r ≤ 0.99) but show negative correlation with canning yield percentage (-0.38 ≤ r ≤ 0.04). The nett yield of abalone shows a relatively strong positive correlation with the live weight (r = 0.66). Shell length is highly heritable (h2 ≈ 0.48) and show a strong positive correlation with live weight (r = 0.94). Shell weight is also highly correlated with live weight (r = 0.80) and the SW: PGW ratio does not show a significant correlate with live weight (r = 0.03). Weight-related traits show heritability values ranging from 0.15 to 0.20 that could allow a positive genetic response. Shell length (as a linear growth parameter) shows a high heritability (h2 ≈ 0.48) and a strong positive correlation with live weight (r = 0.94) which also makes it suitable for use as a selection criterion in breeding programmes for improved growth rate. Direct selection for canning yield is compromised by the destructive nature of measurement and the low heritability (h2 < 0.10). The negative correlations between yield as a percentage and growth traits (-0.38 ≤ r ≤ 0.04) further complicate its use as a direct breeding objective. Although the canning yield as a percentage shows a decrease with an increase in live weight, the nett canning yield increases (r = 0.66) with the live weight. It is therefore recommended to use shell length as a criterion for selection for increased growth rate and nett yield, thereby optimising profitability.
A quantitative genetic analysis of the effect of crossbreeding on the growth rate of the South African abalone, Haliotis midae
(Stellenbosch : Stellenbosch University, 2003-04) Vorster, Gysbert; Brink, Danie; Stellenbosch University. Faculty of Agrisciences. Dept. of Genetics.
ENGLISH ABSTRACT: The genetic status of H. midae broodstock in the South African aquaculture industry reflects that of random samples originating from undomesticated wild populations. The nature of growth in abalone is very slow, taking between three and four years to reach a marketable size of between 60 to 100 grams. It is therefore of paramount importance to improve this trait in order to ensure global competitiveness and economic viability within the industry. Improving this negative characteristic through conventional selection methods is a long-term venture and alternative means that would yield instantaneous results had to be considered. Crossbreeding was identified as an alternative, short-term strategy to improve growth rate. A crossbreeding experiment was performed between two populations of the abalone, Haliotis midae, from the East (E) and West (W) Coast of South Africa. This was done to investigate the occurrence of heterosis for growth among the crossbred genotypes (East x West, West x East). Fifteen males and females from both the East and the West Coast populations were mated in a complete dialelle crossbreeding experiment to produce four progeny groups (WW, EE, EW and WE). Progeny groups were evaluated for weight (bW) and length gain (bL) over a specific growth period of 9 months. The results provided no evidence of significant differences in weight (P = 0.085) or length gain (P = 0.244) between the four progeny groups, giving no indication of significant heterosis for weight and length gain among the crossbred progenies of these East and West Coast populations. It is recommended that further efforts to obtain improved growth rate in the abalone, Haliotis midae, through crossbreeding only be considered in light of clear evidence of substantial genetic differentiation between such populations.
Variation in sex determination and the application of the YY male technology for the production of all-male populations of the tilapia Oreochromis mossambicus
(Stellenbosch : Stellenbosch University, 2004-12) Jordaan, M. S.; Brink, Danie; Stellenbosch University. Faculty of Agrisciences. Dept. of Genetics.
ENGLISH ABSTRACT: Tilapia is presumed to have a well-defined genetic mechanism of sex determination, but not all sex ratios are compatible with a monofactorial sex determination model. A theory of autosomal gene influence, as well as temperature sex determination (TSD) has been proposed in order to explain large variations in sex ratios. This study assessed the variation in progeny sex ratio in O. mossambicus as a basis for the application of YY male technology in the production of all-male progeny groups. Three populations of O. mossambicus were sampled that are representative of the geographical distribution of the species in Southern Africa. Progeny groups were produced from randomly selected parents and maintained at constant temperature during the labile period of sex differentiation. Variation in sex ratios between different families of the same strain as well as between different strains was calculated. No significant differences were observed in sex ratio between strains, though a significant intra-group variation was identified. This study identified both male and female-biased sex ratios. The data in general conform to a monofactorial sex determination model. Male-biased sex ratio observed in one strain can possibly be ascribed to temperature sex determination (TSD). This strain of O. mossambicus originated from an area with different annual temperature patterns and the possibility of TSD having an adaptive advantage is discussed. This thesis also presents the results of a program to produce monosex male tilapia through the application of the YY male technology in O. mossambicus. Viable XY female and YY male genotypes were produced. XY females sired progenies ranging from 68-94% male ofspring, while YY males sired a mean progeny of 94% male phenotypes. From these results it is concluded that YY male technology provide a viable method for the production of all-male progeny in O. mossambicus. Once available on a commercial scale, the technology can be made more reliable through the application of the appropriate selection methods.

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