Genes and growth performance in crustacean species : a review of relevant genomic studies in crustaceans and other taxa

Global aquaculture has expanded rapidly to address the increasing demand for aquatic protein needs and an uncertain future for wild fisheries. To date, however, most farmed aquatic stocks are essentially wild and little is known about their genomes or the genes that affect important economic traits in culture. Biologists have recognized that recent technological advances including next generation sequencing (NGS) have opened up the possibility of generating genome wide sequence data sets rapidly from non-model organisms at a reasonable cost. In an era when virtually any study organism can 'go genomic', understanding gene function and genetic effects on expressed quantitative trait locus phenotypes will be fundamental to future knowledge development. Many factors can influence the individual growth rate in target species but of particular importance in agriculture and aquaculture will be the identification and characterization of the specific gene loci that contribute important phenotypic variation to growth because the information can be applied to speed up genetic improvement programmes and to increase productivity via marker-assisted selection (MAS). While currently there is only limited genomic information available for any crustacean species, a number of putative candidate genes have been identified or implicated in growth and muscle development in some species. In an effort to stimulate increased research on the identification of growth-related genes in crustacean species, here we review the available information on: (i) associations between genes and growth reported in crustaceans, (ii) growth-related genes involved with moulting, (iii) muscle development and degradation genes involved in moulting, and; (iv) correlations between DNA sequences that have confirmed growth trait effects in farmed animal species used in terrestrial agriculture and related sequences in crustacean species. The information in concert can provide a foundation for increasing the rate at which knowledge about key genes affecting growth traits in crustacean species is gained.

Modeling meat yield based on measurements of body traits in genetically improved giant freshwater prawn (GFP) Macrobrachium rosenbergii

A single-generation dataset consisting of 1,730 records from a selection program for high growth rate in giant freshwater prawn (GFP, Macrobrachium rosenbergii) was used to derive prediction equations for meat weight and meat yield. Models were based on body traits [body weight, total length and abdominal width (AW)] and carcass measurements (tail weight and exoskeleton-off weight). Lengths and width were adjusted for the systematic effects of selection line, male morphotypes and female reproductive status, and for the covariables of age at slaughter within sex and body weight. Body and meat weights adjusted for the same effects (except body weight) were used to calculate meat yield (expressed as percentage of tail weight/body weight and exoskeleton-off weight/body weight). The edible meat weight and yield in this GFP population ranged from 12 to 15 g and 37 to 45 %, respectively. The simple (Pearson) correlation coefficients between body traits (body weight, total length and AW) and meat weight were moderate to very high and positive (0.75–0.94), but the correlations between body traits and meat yield were negative (−0.47 to −0.74). There were strong linear positive relationships between measurements of body traits and meat weight, whereas relationships of body traits with meat yield were moderate and negative. Step-wise multiple regression analysis showed that the best model to predict meat weight included all body traits, with a coefficient of determination (R 2) of 0.99 and a correlation between observed and predicted values of meat weight of 0.99. The corresponding figures for meat yield were 0.91 and 0.95, respectively. Body weight or length was the best predictor of meat weight, explaining 91–94 % of observed variance when it was fitted alone in the model. By contrast, tail width explained a lower proportion (69–82 %) of total variance in the single trait models. It is concluded that in practical breeding programs, improvement of meat weight can be easily made through indirect selection for body trait combinations. The improvement of meat yield, albeit being more difficult, is possible by genetic means, with 91 % of the variation in the trait explained by the body and carcass traits examined in this study.

The introduction of transgenes to control blackheart in pineapple (Ananas Comosus L.) cv. Smooth Cayenne by microprojectile bombardment

A transformation technique for the introduction of transgenes to control blackheart by particle bombardment has been developed for pineapple cv. Smooth Cayenne. Leaf callus cultures capable of high frequency organogenesis with a short regeneration time were used as explant material. Gus and gfp reporter genes were used to observe and determine transient and stable expression. The ppo gene, isolated from pineapple, was introduced to control blackheart. Co-transformation occurred with constructs containing the nptII gene conferring geneticin resistance. We have recovered 15 independent transgenic gus and gfp lines each from 8 separate experiments and 22 ppo lines from 11 experiments. Gus, gfp, ppo and nptII positive plants have been regenerated, which have been shown by Southern blot analysis to be stable transgenics containing multiple copies of the introduced genes. These results show that biolistic gene delivery in pineapple can be successfully achieved at an acceptable efficiency of 0.21-1.5% for genetic improvement of 'Smooth Cayenne', the industry standard throughout the world.

The domestication of African mahogany (Khaya senegalensis) in northern Australia.

African mahogany has demonstrated much potential, in many field trials spanning several decades and in furniture manufacturers' evaluations, as a high-value timber species for plantations in northern Australia. It is in the early stages of domestication via a low-intensity, informally-collaborative, mostly-public-sector program of conservation and genetic improvement begun 5 y ago. Silvicultural techniques are being developed through experience in both 'small grower' and larger-scale plantings. See this issue's cover for photographs.

Screening selected myrtaceous tree species for production of essential oils in northern Queensland

Trials to identify alternative cropping options to Melaleuca alternifolia for northern Queensland essential oil growers were established at Dimbulah and Innot Hot Springs in 2001. Seed sources of Asteromyrtus symphyocarpa (1,8-cineole form), Eucalyptus staigeriana (citral), Melaleuca cajuputi subsp. cajuputi (trans-nerolidol), M. ericifolia (d-linalool), M. quinquenervia (trans-nerolidol and viridiflorol forms) and M. viridiflora (methyl cinnamate) with potential to produce commercial foliar oils were evaluated. Information was gathered on their adaptability, growth and oil yields over 49 months and 52 months (two harvests) from planting at Dimbulah and Innot Hot Springs, respectively. Of the species and chemotypes evaluated, M. quinquenervia showed potential for commercial production of trans-nerolidol, a compound used in perfumery. It had a very high survival rate (96%) and yields could be expected to improve dramatically from the average 100 kg/ha per harvest achieved in these trials with further research into selection of seed source, control of insect damage and breeding for genetic improvement. M. cajuputi subsp. cajuputi gave a similar performance to M. quinquenervia. The rarity of the trans-nerolidol form of this species and remoteness of its natural occurrence are impediments to further planting and research. E. staigeriana, with second harvest yields of ~600 kg/ha, performed exceptionally well on both sites but potential for development is limited by the ready availability of competitively priced E. staigeriana oil produced in South America. Survival of M. ericifolia ranged from 62% to 82% at 32 months (second harvest) at Innot Hot Springs and was deemed a failure at Dimbulah with poor growth and low survival, raising a major question about the suitability of this species for cultivation in the seasonally dry tropics. Planting of this species on a wider scale in northern Queensland cannot be recommended until more is known about factors affecting its survival. A. symphyocarpa and M. viridiflora were too slow-growing to warrant further consideration as potential oil-producing species at this time.

Optimising between and within family selection for harvest weight in prawns with restricted harvest size.

Manual grading of prawns restricts the number that can be harvested. A restricted harvest size places a limit on the opposing within family and between family sources of selection pressure. A simulation study with inbreeding constrained at 0.5% per generation, a harvest size of 2000, heritability of 0.3, common family environmental effect of 0.1, indicates that maximum response to selection could be achieved with as few as 40 families. Increasing the number of families above 80 may reduce total selection response. It is important to be aware that increasing the number of families may not always yield a greater genetic response.

Reproductive Biology of Corymbia citriodora subsp. variegata and Effective Pollination across its Native Range.

The spotted gum species complex represents a group of four eucalypt hardwood taxa that have a native range that spans the east coast of Australia, with a morphological cline from Victoria to northern Queensland. Of this group, Corymbia citriodora subsp. variegata (CCV) is widespread in south-eastern Queensland and northern New South Wales. It is currently the most commonly harvested native hardwood in Queensland. However, little basic knowledge of the reproductive biology of the species is available to inform genetic improvement and resource management programmes. Here we take an integrative approach, using both field and molecular data, to identify ecological factors important to mating patterns in native populations of CCV. Field observation of pollinator visitation and flowering phenology of 20 trees showed that foraging behaviour of pollinator guilds varies depending on flowering phenology and canopy structure. A positive effect of tree mean flowering effort was found on insect visitation, while bat visitation was predicted by tree height and by the number of trees simultaneously bearing flowers. Moreover, introduced honeybees were observed frequently, performing 73% of detected flower visits. Conversely, nectar-feeding birds and mammals were observed sporadically with lorikeets and honeyeaters each contributing to 11% of visits. Fruit bats, represented solely by the grey-headed flying fox, performed less than 2% of visits. Genotyping at six microsatellite markers in 301 seeds from 17 families sampled from four of Queensland's native forests showed that CCV displays a mixed-mating system that is mostly outcrossing (tm = 0.899 ± 0.021). Preferential effective pollination from near-neighbours was detected by means of maximum-likelihood paternity analysis with up to 16% of reproduction events resulting from selfing. Forty to 48% of fertilising pollen was also carried from longer distance (>60 m). Marked differences in foraging behaviour and visitation frequency between observed pollinator guilds suggests that the observed dichotomy of effective pollen movement in spotted gums may be due to frequent visit from introduced honeybees favouring geitonogamy and sporadic visits from honeyeaters and fruit bats resulting in potential long-distance pollinations.

Transgenic rootstocks for production of GM-free fruit

Development of regeneration and transformation methods for genetic improvement of rootstocks for mango, avocado and citrus.

Understanding the cropping behaviour of Riberry (Syzygium leuhmannii)

This study highlights the complexity of flowering biology in Syzygium and demonstrates how a basic understanding of a species’ fundamental biology is necessary for successful commercial cultivation. This report brings together useful information from previous international research on Syzygium as well as providing a basic understanding of flower biology, the nature of fruit set and seediness in riberry. Much of these findings have implications for the cultural management of riberry orchards to optimise fruit set and minimise seed set. It raises the possibility of avenues for genetic improvement.

Lessons from established breeding programs: terrestrial and aquatic animals

Some relevant components of selection program theory and implementation are reviewed. This includes pedigree recording, genetic evaluation, balancing genetic gains and genetic diversity and tactical integration of key issues. Lessons learned are briefly described – illustrating how existing method and tools can be useful when launching a program in a novel species, and yet highlighting the importance of proper understanding and custom application according to the biology and environments of that species.

Breeding programs on Atlantic salmon in Norway: lessons learned

An early establishment of selective breeding programs on Atlantic salmon has been crucial for the success of developing efficient and sustainable salmon farming in Norway. A national selective breeding program was initiated by AKVAFORSK at the beginning of the 1970s, by collecting fertilized eggs from more than 40 Norwegian river populations. Several private selective breeding programs were also initiated in the 1970s and 1980s. While these private programs were initiated using individual selection (i.e. massselection) to genetically improve growth, the national program was designed to gradually include all economically important traits in the breeding objective (i.e. growth, age at sexual maturation, disease resistance and quality traits) using a combined family and within-family selection strategy. Independent of which selection strategy and program design used, it is important to secure and maintain a broad genetic variation in the breeding populations to maximize selection response. It has been documented that genetically improved salmon from the national selective breeding program grow twice as fast as wild Atlantic salmon and require 25 per cent less feed, while salmon representing the private breeding programs all show an intermediate growth performance. As a result of efficient dissemination of genetically improved Atlantic salmon, the Norwegian salmon farming industry has reduced its feed costs by more than US$ 230 million per year! The national selective breeding program on Atlantic salmon was commercialized into a breeding company (AquaGen) in 1992. Five years later, several private companies and the AKVAFORSK Genetics Center (AFGC) established a second breeding company (SalmoBreed) using breeding candidates from one of the private breeding programs. These two breeding companies have similar products, but different strategies on how to organize the breeding program and to disseminate the genetically improved seed to the Norwegian salmon industry. Greater competition has increased the necessity to document the genetic gain obtained from the different programs and to market the economic benefits of farming the genetically improved breeds. Both breeding companies have organized their dissemination to get a sufficient share of the economic benefits in order to sustain and improve their breeding programs.

Lessons from the breeding program on common carp in Hungary

Common carp is one of the most important cultured freshwater fish species in the world. Its production in freshwater areas is the second largest in Europe after rainbow trout. Common carp production in Europe was 146,845 t in 2004 (FAO Fishstat Plus 2006). Common carp production is concentrated mainly in Central and Eastern Europe. In Hungary, common carp has been traditionally cultured in earthen ponds since the late 19th century, following the sharp drop in catches from natural waters, due to the regulation of main river systems. Different production technologies and unintentional selection methods resulted in a wide variety of this species. Just before the intensification of rearing technology and the exchange of stocking materials among fish farms (early sixties), “landraces” of carp were collected from practically all Hungarian fish farms into a live gene bank at the Research Institute for Fisheries, Aquaculture and Irrigation (HAKI) at Szarvas (Bakos and Gorda 1995; Bakos and Gorda 2001). In order to provide highly productive hybrids for production purposes starting from 1964, different strains and crosses between Hungarian landraces were created and tested. During the last 40 years, approximately 150 two-, three-, and four-line hybrids were produced. While developing parental lines, methods of individual selection, inbreeding, backcrossing of lines, gynogenesis and sex reversal were used. This breeding program resulted in three outstanding hybrids: “Szarvas 215 mirror” and “Szarvas P31 scaly” for pond production, and “Szarvas P34 scaly” for angling waters. Besides satisfying the needs of industry, the live gene bank helped to conserve the biological diversity of Hungarian carp landraces. Fifteen Hungarian carp landraces are still maintained today in the gene bank. Through exchange programs fifteen foreign carp strains were added to the collection from Central and Eastern Europe, as well as Southeast Asia (Bakos and Gorda 2001). Besides developing the methodology to maintain live specimens in the gene bank, the National Carp Breeding Program has been initiated in cooperation with all the key stakeholders in Hungary, namely the National Association of Fish Producers (HOSZ), the National Institute for Agricultural Quality Control (OMMI), and the Research Institute for Fisheries, Aquaculture and Irrigation (HAKI). In addition, methodologies or technologies for broodstock management and carp performance testing have been developed. This National Carp Breeding Program is being implemented successfully since the mid-1990s.

Lessons from the breeding program of rohu

For the first time in India, selective breeding work has been initiated at the Central Institute of Freshwater Aquaculture, Bhubaneswar, India in collaboration with the Institute of Aquaculture Research (AKVAFORSK), Norway. Rohu has been chosen as the model species because it enjoys the highest consumer preference among Indian major carps (IMC) although its performance was observed to be slower than other IMC. As this was the first ever selection work on any Indian major carp, many procedures and techniques for successful implementation of the programs were standardized (i.e. production of full-sib groups, establishment of model hatchery for selective breeding of carps, rearing of full-sib groups in partitioned nursery ponds, individual tagging with the Passive Integrated Transponder (PIT) tag, communal rearing, sampling, data analysis, field testing and dissemination of improved rohu). After four generations of selection, an average of 17 per cent higher growth per generation was observed in improved rohu.

Policies on release of improved fish strains in China

Since 1991, the certification, release and maintenance of new species for aquaculture have become part of the national policy in China. During the past 15 years, this policy has been conducted and improved and has begun to show its significant role in Chinese fisheries. This paper describes the updated system of certification, release and maintenance of new species for aquaculture in China.

Maintaining the "public good" nature of improved fish strains: dissemination of knowledge and materials

Many sources of information that discuss currents problems of food security point to the importance of farmed fish as an ideal food source that can be grown by poor farmers, (Asian Development Bank 2004). Furthermore, the development of improved strains of fish suitable for low-input aquaculture such as Tilapia, has demonstrated the feasibility of an approach that combines “cutting edge science” with accessible technology, as a means for improving the nutrition and livelihoods of both the urban poor and poor farmers in developing countries (Mair et al. 2002). However, the use of improved strains of fish as a means of reducing hunger and improving livelihoods has proved to be difficult to sustain, especially as a public good, when external (development) funding sources devoted to this area are minimal1. In addition, the more complicated problem of delivery of an aquaculture system, not just improved fish strains and the technology, can present difficulties and may go explicitly unrecognized (from Sissel Rogne, as cited by Silje Rem 2002). Thus, the involvement of private partners has featured prominently in the strategy for transferring to the public technology related to improved Tilapia strains. Partnering with the private sector in delivery schemes to the poor should take into account both the public goods aspect and the requirement that the traits selected for breeding “improved” strains meet the actual needs of the resource poor farmer. Other dissemination approaches involving the public sector may require a large investment in capacity building. However, the use of public sector institutions as delivery agents encourages the maintaining of the “public good” nature of the products.