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Traditionally, male pigs are surgically castrated without anaesthesia to avoid later occurrence of the sex odour of androstenone in the carcass. Active immunization against gonadotropin-releasing hormone (GnRH) is a painless alternative which inhibits LH and thus steroidogenesis in the Leydig cells. In a preceding study we clarified the return of Leydig cell function after the last dose of antigen by measuring hormones, and found a considerable variation (10 to 24 weeks) till return of their function (testosterone ⩾ 0.5 ng/ml blood plasma). The present paper analyses histological data on testes characteristics of the same six boars at an age of 52 weeks (26 weeks after last immunization). Data were compared to another four boars which were not immunized but slaughtered at the same age. Testis weight was related to the concentration of testosterone in blood. In boars, that first returned to testicular function, testis weight even exceeded those in controls probably due to rebound phenomena. Differences in testis weight were mainly due to differences of Leydig cell content of cytoplasm, and less to the size of nuclei. Additionally, the height of seminiferous epithelium was slightly dependent on testosterone concentrations and contributed moderately to differences in testis weight. Altogether, normalization of testicular function, even after return to steroidogenesis, requires another 13 weeks.
A central goal of conservation is the maintenance of ecosystems, species or populations at their current state, or the restoration of biological systems to some former state. In cases of recent ecological collapse, such as the decline of a population due to the introduction of disease or an invasive competitor, or due to over-hunting or habitat destruction, we may have monitored the process from start to finish and the former state may be sufficiently well-described to give us a target for restoration. In most cases, though, serious monitoring only begins after a decline is identified, and only anecdotal evidence is available to guide our reconstruction of the past. In addition, the mechanisms that have driven changes in ecosystems are typically unknown. Are these changes the result of natural processes acting over many millennia, or has human activity drastically altered the natural trajectory? The recent application of genetics to conservation has allowed us to describe more fully the current status of populations by quantifying such properties as levels of inbreeding, effective population sizes, levels of genetic variation, and gene flow (Fleischer 1998; DeSalle and Amato 2004). Through the application of coalescent models, population genetics has also given us insight into the historical status of populations, whether such properties as size and growth of a population have changed and on what time scale these changes have occurred. Unfortunately, the stochastic nature of the coalescent process and the effects of selection often impair our ability to confidently reconstruct historical states. With the relatively recent development of ancient DNA (aDNA) techniques, however, we can now step directly backwards in time to characterize historical genetic diversity and to better understand the processes that have generated current levels of genetic diversity and population structure. Our ability to travel back in time using aDNA has allowed us to view conservation issues with a broader temporal perspective and has provided a better framework for understanding the impact of humans in shaping contemporary animal populations.
A key component of the emergence of conservation genetics as a recognisable subdiscipline of conservation biology over the last ten years has been the development of methods to genetically assess and monitor populations of endangered species non-invasively. The rapid development of methodologies for ensuring the accurate capture of molecular data from elusive, easily stressed or potentially dangerous (!) organisms and concerns over the accuracy of the data produced have prompted a number of excellent reviews on the subject in recent times (e.g. Taberlet et al. 1999; Taberlet and Luikart 1999; Piggott and Taylor 2003; Woodruff 2003; Wayne and Morin 2004). Here, we will review the issues and wide-ranging applications of non-invasive genetic analysis without focusing on the molecular technicalities in great detail.
Why non-invasive genetics?
Before the advent of non-invasive genetics and the use of animal by-products such as faeces, shed hair, feathers, bones, fish scales, teeth, etc., obtaining genetic material from wild populations was often ethically (in particular for species listed as endangered and critically endangered under CITES regulations) and logistically extremely difficult. Now such analysis is increasingly possible and the sampling of large populations without visual/physical contact is particularly beneficial for endangered species or if the species studied can transmit or are susceptible to disease (e.g. great ape species whose pathogens are often extremely similar to those of the researchers studying them). In a century where linking behaviour, social structure, dispersal and population genetic structure has become a new challenge for conservation geneticists, the development of non-invasive sampling and genotyping has provided the opportunity to explore these links and has dramatically opened new areas for research.
The characterisation of non-invasive material using molecular markers allows biologists to identify and count individuals in wild populations, identify the sex of those individuals and determine their movement patterns, infer parentage, kinship and relatedness, and assess pathogens and diet (see Kohn and Wayne 1997 for a review).
Iberian pigs and wild boars are the source of highly priced meat and dry-cured products. Iberian maternal origin is mandatory for labeled Iberian products, making necessary the authentication of their maternal breed origin. Discrimination between wild and domestic pig maternal origin may be useful to distinguish labeled wild boar meat obtained from hunting or farming. In order to detect useful polymorphisms to trace Iberian, Duroc and wild boar maternal lineages, we herein investigated the complete porcine mitochondrial DNA (mtDNA) using three complementary approaches. Near-complete mtDNA sequences (16989 bp), excluding the minisatellite present in the displacement loop region (D-loop), were successfully determined in six Iberian pigs, two Duroc and six European wild boars. To complete the mtDNA analysis, the D-loop minisatellite region was also analyzed in the same set of samples by amplification and capillary electrophoresis detection. Finally, the frequencies of Asian and European Cytochrome B (Cyt B) haplotypes were estimated in Iberian (n = 96) and Duroc (n = 125) breeds. Comparison of near-complete mtDNA sequences revealed a total of 57 substitutions and two Indels. Out of them, 32 polymorphisms were potential Iberian markers, 10 potential Duroc markers and 16 potential wild boar markers. Fourteen potential markers (five Iberian and nine Duroc), were selected to be genotyped in 96 Iberian and 91 Duroc samples. Five wild boar potential markers were selected and tested in samples of wild boars (73) and domestic pigs including: 96 Iberian, 16 Duroc, 16 Large White and 16 Landrace. Genotyping results showed three linked markers (m.7998C>T, m.9111T>C, m.14719A>G) absent in Duroc and present in Iberian pigs with a frequency 0.72. Six markers (m.8158C>T, m.8297T>C, m.9230G>A, m.11859A>G, m.13955T>C, m.16933T>C), three of them linked, were absent in Iberian pigs and present in Duroc with a joint frequency of almost 0.50. Finally three linked markers (m.7188G>A, m.9224T>C, m.15823A>G) were solely detected in wild boars with a frequency 0.22. The D-loop minisatellite results showed overlapping ranges of fragment sizes and suggested heteroplasmy, a result that nullify the use of this region for the development of breed diagnostic markers. The Cyt B haplotype results showed the presence of European haplotypes in Iberian while one of the Asian haplotypes was detected in Duroc with a frequency 0.22, linked to the Duroc marker m.9230G>A. Our results are valuable to resolve the problems of Iberian and wild boar maternal origin determination but additional markers are required to achieve totally useful genetic tests.
A study was done to evaluate the effect of using progesterone (P4) intravaginal device (CIDR: controlled internal drug-releasing dispenser) to synchronise the return to oestrus of previously timed inseminated (TAI) dairy heifers, and to evaluate embryo survival and pregnancy rate (PR) in the return to oestrus heifers. At the onset of the artificial insemination (AI) breeding period (day −9), heifers were randomly assigned into two groups (treated group CGPG, n = 79) and (control group GPG, n = 83). Every heifer in both groups was injected with gonadotropin-releasing hormone (GnRH) agonist and prostaglandin F2-alpha (PGF2α) as follows: GnRH on day −9; PGF2α on day −2; GnRH and TAI on day 0. Heifers in both groups received TAI within 30 min after the second GnRH injection. Artificial insemination at first breeding was conducted for all heifers during 55 days from day 0. On day 14 after timed insemination, every heifer in the CGPG group received CIDR device for 6 days. Within 3 days after CIDR removal, more heifers in CGPG group showed oestrus within 1.9 days compared to heifers that showed oestrus within 2.9 days in the control. Within 10 days after CIDR removal, more heifers in the CGPG group showed oestrus within 2.4 days compared to heifers that showed oestrus within 6.7 days in the control. PRs on days 30 and 55 were not different between both groups, while PR on day 55 during September were higher (P = 0.032) in CGPG group (58.0%) than GPG group (37.0%). In addition, PR from first to second AI was higher (P = 0.037) for CGPG group (79.8%) than for GPG group (65.1%) but it was similar after that. Pregnancy losses between days 30 and 55 tended to be lower (P = 0.089) for the CGPG group (12.7%) compared to 25.1% for the GPG group. Interval between first and second AI was lower (P = 0.052) for the CGPG group (27.5 ± 1.6 days) compared to 31.6 ± 1.3 days for heifers in the GPG group but no differences were detected for intervals from second to third AI and from third to fourth AI between the two groups. Number of services per pregnancy was not different between CGPG and GPG groups. Results indicate that the CIDR device improved synchronisation to return to oestrus and increased PR to first AI during high temperature months by reducing embryonic losses.
When it comes to advocating animal conservation, it is difficult to be convincing without becoming alarmist. The fact is, time is running out for many of the world's animal species. Habitat loss, introduced species, overexploitation and pollution, all caused by human activities, combine with stochastic factors to place ever-increasing pressure on natural populations (Primack 2002). The estimates of the mid-1990s, predicting that thousands of species and millions of unique populations would go extinct in the following decades (Ehrlich and Wilson 1991; Smith et al. 1993; Lawton and May 1995), remain as relevant as when they were first made, and we are still living in an era of unprecedented biodiversity loss, with current extinction rates 100–1000 times the background rate (Primack 2002) and 5000–25 000 times that recorded in the fossil record (Frankham et al. 2002; but see Mace et al. 1996). Recently, however, there have been some positive signs in the media that biologists’ warnings are being received (e.g. Gianni 2004; Devine et al. 2006; Black 2006; Gabriel 2007; Stern 2007), and a rapid and efficient approach in providing information pertinent to biodiversity preservation could be pivotal in policy decision and in optimizing resource allocation (Naidoo and Ricketts 2006; Marsh et al. 2007). Since the foundation of the field of conservation biology, it has been argued that a synergy between conservation biology and advanced population genetics could provide important information that policy-makers need. As should be obvious by its title, the purpose of this book is an attempt to go some way towards maturing such a synergy; hence, this introduction presents a brief history and the current state of this partnership.
THE EXTINCTION CRISIS
In order to be convinced of the urgency for animal conservation and the information necessary to practise it, an update on the current extinction crisis is pertinent.
This paper constitutes a review on the welfare aspects of piglet castration that considers the scientific literature published after 2004. Castrating during the neonatal period (1 to 3 days of age) is clearly painful. In addition, inflammatory processes may take place at the sites of incision, thus adding further pain to the procedure. Surgical castration with general and local anaesthesia, in combination with long-term analgesia, has been shown to reduce pain but the additional handling and injection of the anaesthetic, the effectiveness and limited safety margins have to be thoroughly evaluated. Raising entire males during the whole fattening period or immunocastration of males towards the end of the fattening period are other alternatives with welfare benefits in young pigs compared to current surgical castration, but with some potential welfare drawbacks regarding handling stress and behaviour during fattening. Based on the current knowledge, it can be concluded that sperm sexing and raising entire males after genetic control of boar taint are potentially preferable alternatives to current practices, but need further research, as these methods are not yet available.
In order to investigate the effects of short road transport stress on total and free iodothyronines, body weight (BW), rectal temperature and heart rate (HR) changes, 126 healthy stallions were studied in basal conditions, before and after transport. A total of 60 Thoroughbred and 66 crossbred stallions aged 4 to 15 years with previous travelling experience were transported by road in a commercial trailer for a period of about 3 to 4 h (distance under 300 km). Blood samples and functional variables were collected in each horse box, one week before loading and transport in basal conditions (control samples), one week later immediately before loading (pre-samples) and again after transport and unloading (about 3 to 4 h) in each new horse box, within 30 min of their arrival at the breeding stations (post-samples). Compared to the before-transport values, increases in circulating T3, T4 and fT4 levels (P < 0.01) were observed after transport, irrespective of breed, but not for fT3 levels. Lower T4 and fT4 levels were observed in basal II (at 1100 h) (P < 0.01) than in basal I (at 0800 h) conditions and before transport. Thoroughbreds showed higher fT3 (P < 0.05) and fT4 (P < 0.01) levels after transport than crossbred stallions. No significant differences were observed for T3 and T4. Compared to the before-transport values, significant increases in rectal temperature (P < 0.01) and HR (P < 0.05) were observed after transport. No differences were observed between basal I, II and before values for functional variables. Significant correlations between T3 and rectal temperature, BW and HR were found. The results indicate that short road transport induces a preferential release of T3, T4 and fT4 hormones from the thyroid gland in relation to different breed, and an increase in rectal temperature and HR. No significant changes in BW were observed. No differences were observed in relation to different ages. The data obtained suggest that the stallion’s thyroid hormones and functional variables may play an important role in assessing the effects of transport stress and a horse’s coping strategy.
Tibetan Mastiff is one of the most archaic, ferocious and the largest dogs in the world. A total of 140 individuals from four geographically separated populations in China (Tibet, Gansu, Qinghai and Beijing) were sampled and genetic diversity was assessed using 10 microsatellite loci on eight different chromosomes. The mean number of alleles per locus ranged from 6 to 13. The mean observed and expected heterozygosities, polymorphism information content and allelic richness were 0.69, 0.79, 0.76 and 7.59, respectively, indicating relatively high genetic diversity in Tibetan Mastiff. However, a highly significant deficiency in heterozygote was observed within populations (mean FIS = 0.11, bootstrap 95% confidence interval (0.06, 0.17)) and total inbreeding (mean FIT = 0.12, bootstrap 95% confidence interval (0.06, 0.18)), along with strong inbreeding coefficients within populations (Fis > 0.09), all of which suggested that intense inbreeding practices occurred in Tibetan Mastiff. Therefore, effective and appropriate breeding management projects in present Tibetan Mastiff will be desirable and urgent. Low genetic differentiation was obtained with a mean FST of 0.01 (bootstrap 95% confidence interval (0.007, 0.019)). Additionally, the four Tibetan Mastiff populations showed close relationships in the neighbor-joining polygenetic tree based on the coancestral genetic distances. Tibetan Mastiff was investigated by using microsatellite loci at the first time, which could facilitate the better understanding of present situation at the molecular level, breed conservation and utilization in Tibetan Mastiff.
A standardised methodology is required for classification of starchy diets. One important question is what feeding status is optimal to describe glycaemic and insulinaemic responses to the respective starchy diet. Four horses were fed, in a randomised order, four different feeding protocols relative to offering hay before or after cracked corn (CC): (i) ad libitum hay for 12 h before feeding CC and ad libitum hay after CC intake for 9 h (+CC+), (ii) ad libitum hay for 12 h before feeding CC and no hay after CC intake for 9 h (+CC−), (iii) feed restriction for 12 h before feeding CC and ad libitum hay after CC intake for 9 h (−CC+) and (iv) 1.2 kg hay/100 kg body weight (BW) per day, divided into two equal portions and offered at 0900 h and 1900 h, feed restriction for 12 h before feeding CC and no hay after CC intake for 9 h (−CC−). CC intake was adjusted to a starch intake of 2 g/kg BW. The different hay offerings did not affect basal plasma glucose and insulin levels. A significant rise in plasma glucose and insulin was found after CC intake for all diets. The highest peak glucose levels were analysed for −CC+, and the lowest glucose peaks were found for +CC− (diet P < 0.05). The highest insulin peaks were monitored for −CC+ (31.27 ± 18.19 μU/ml) and lower peaks for +CC− (13.36 ± 2.93 μU/ml) (diet P < 0.05). Insulin for −CC− and +CC− returned to resting values about 300 min after CC feeding. For +CC+ and −CC+, insulin levels were still above resting levels 510 min after CC intake (diet P < 0.05). The present data suggest that feed restriction for 12 h before feeding the starchy diet and no further roughage intake during blood sampling period provide the best-defined conditions.
Group-living animals travel together to collectively exploit the resources of their environment. This study investigates how social relationships and individual temperament traits affect movement orders in domestic cattle and sheep. We analysed spontaneous group movements occurring at pasture after a resting period in a group of 15 18-month-old Charolais heifers and a group of 19 1-year-old Romane ewe-lambs. For each species, animals had similar social experience and no kinship ties. Before that, animals were observed within the group to establish their social status (e.g. dominance and preferential relationships, and sociability), then in individual tests in order to assess their emotional traits. In both species, most individuals could initiate a group movement but some individuals were more successful than others in recruiting the rest of the group. Ewe-lambs, and to a lesser extent heifers, held preferential positions during travel. We did not find any significant correlations in either species between animal order and their position in the dominance hierarchy (heifers: P = 0.438; ewe-lambs: P = 0.574) while individuals linked by preferential bonds frequently followed each other during group movements (heifers: P < 0.001; ewe-lambs: P < 0.001). With regard to social traits, heifers with a low cohesion index, and with a lower number of partners with whom they develop frequent affinitive interactions, acted more frequently as ‘first movers’ (P = 0.040 and 0.023, respectively), as well as did ewe-lambs with a high spatial independency index (P = 0.002). Ewe-lambs with the highest cohesion indices were more frequently observed in front of the group while moving halfway between departure and arrival (P = 0.028). We did not find significant correlations between individual positions during group movements and emotional traits such as reactivity, boldness and fearfulness. We conclude that preferential bonds and individual traits related to social dependence were more influential in spontaneous group movements at pasture than were emotional traits and dominance status.
This study investigates the effects of two feeding systems and two dairy cow breeds on milk yield and composition, physical and sensorial properties of Camembert and Pont-l’Evêque cheeses. The experiment consisted of a 2 × 2 factorial arrangement of treatments. A low energy grass diet with only 15% of concentrate (LowGS) was compared with a high-energy maize silage diet with 30% concentrate (HighMS). Thirty-four Holstein (Ho) and 34 Normande (No) cows in early lactation were assigned to one of two feeding systems for a 6-week period. Cows on the LowGS feeding system had lower milk yield, fat and protein content. In both feeding systems, No cows had lower milk yields but higher milk protein contents than Ho cows. The LowGS feeding system altered milk fatty acid (FA) composition by reducing saturated FA. Breed had only a small impact on milk FA. Concerning milk coagulating properties, only the firmness was reduced by the LowGS feeding and the Ho breed. The effects of breed and feeding system on the protein content of cheeses were more marked in Camembert cheese than in Pont-l’Evêque cheese. However, the Camembert cheese from Ho-LowGS was, in fact, characterized especially by lower protein content. LowGS feeding system and No breed produced more yellow cheeses. Feeding systems had limited effects on the firmness of Camembert and Pont-l’Evêque cheeses measured by penetrometry. In sensory analysis, Ho breed and LowGS feeding produced a Camembert cheese with a more melting texture in the mouth due to the increase of spreadability index and of proteolysis. The type of cheese also had an influence: the effects were more important on Camembert cheese than on Pont-l’Evêque cheese. Only the Ho-LowGS treatment produced a very specific Camembert cheese different from the others. The feeding systems and breed of dairy cow have no determinant effect on PDO (protected designation of origin) Camembert and Pont-l’Evêque cheeses, especially regarding taste. In this kind of trial, despite the effects of feeding systems and breed on milk composition, the role of cheese ripening and microbiology appears to be of considerable importance.
Livestock contribute directly (i.e. as methane and nitrous oxide (N2O)) to about 9% of global anthropogenic greenhouse gas (GHG) emissions and around 3% of UK emissions. If all parts of the livestock production lifecycle are included (fossil fuels used to produce mineral fertilizers used in feed production and N2O emissions from fertilizer use; methane release from the breakdown of fertilizers and from animal manure; land-use changes for feed production and for grazing; land degradation; fossil fuel use during feed and animal production; fossil fuel use in production and transport of processed and refrigerated animal products), livestock are estimated to account for 18% of global anthropogenic emissions, but less than 8% in the UK. In terms of GHG emissions per unit of livestock product, monogastric livestock are more efficient than ruminants; thus in the UK, while sheep and cattle accounted for 32% of meat production in 2006, they accounted for ∼48% of GHG emissions associated with meat production. More efficient management of grazing lands and of manure can have a direct impact in decreasing emissions. Improving efficiency of livestock production through better breeding, health interventions or improving fertility can also decrease GHG emissions through decreasing the number of livestock required per unit product. Increasing the energy density of the diet has a dual effect, decreasing both direct emissions and the numbers of livestock per unit product, but, as the demands for food increase in response to increasing human population and a better diet in some developing countries, there is increasing competition for land for food v. energy-dense feed crops. Recalculating efficiencies of energy and protein production on the basis of human-edible food produced per unit of human-edible feed consumed gave higher efficiencies for ruminants than for monogastric animals. The policy community thus have difficult decisions to make in balancing the negative contribution of livestock to the environment against the positive benefit in terms of food security. The animal science community have a responsibility to provide an evidence base which is objective and holistic with respect to these two competing challenges.
The objective of this study was to determine the effect of lamb genotype and plane of nutrition on post-weaning lamb growth, and carcass characteristics in hill sheep systems. The lamb genotypes compared were as follows: purebred Scottish Blackface, Swaledale × Scottish Blackface, Cheviot × Scottish Blackface, Lleyn × Scottish Blackface and Texel × Scottish Blackface lambs. During the finishing period, lambs were allocated to a (i) low plane of nutrition (LP), based on a fixed allowance of dried grass pellets representing a grass-based system; (ii) high plane of nutrition (HP-cereal), with a diet consisting proportionately of 0.20 dried grass pellets : 0.80 cereal-based concentrate offered ad libitum; (iii) high plane of nutrition (HP-cereal/oil), with a diet consisting proportionately of 0.20 dried grass pellets : 0.80 cereal-based concentrate containing oilseed rape offered ad libitum. There were significant genotype × diet interactions for dry matter (DM) intake (P < 0.001), live weight gain (P < 0.05) and carcass gain (P < 0.05). On the LP diet, carcass gains were higher for Texel cross compared to Blackface and Swaledale cross lambs (P < 0.05). On both HP diets, carcass gains were lower in Blackface and Swaledale crosses compared with the Cheviot, Lleyn and Texel cross lambs (P < 0.01). Feed conversion efficiency, expressed in terms of carcass gain per unit of DM, was highest for Texel cross and Cheviot cross, lowest in Swaledale cross and Blackface and intermediate for Lleyn cross lambs (P < 0.001). Conformation classification was lowest for Swaledale cross (P < 0.05) and highest for Texel cross lambs (P < 0.001), whilst those of Blackface, Cheviot and Lleyn cross lambs were similar. Carcasses from Blackface and Lleyn cross lambs had higher fat classifications than Texel and Cheviot cross lambs (P < 0.01). Feed conversion efficiency was lowest with the LP diet compared with the HP diets, resulting in a 5-week longer finishing period and lower carcass weights, fat scores, dressing proportions and subcutaneous fat depth measurements. These results indicate that crossbreeding can have a major effect on lamb performance in the hill sheep sector. Feeding a HP increased lamb growth rates, particularly for high growth-potential genotypes, improved feed efficiency, shortened the finishing period but increased carcass fatness. Including oilseed rape in concentrate-based diets did not affect performance.
Mendel's decision to select such unit characters that would provide insight into the laws of hereditary transmission, whatever their physiological or developmental properties may be, was crucial in the establishment of genetics as an independent discipline of inheritance (Falk, 1995). But when de Vries in 1900 recognized Mendel's work it was because he believed it to contribute to his hypothesis of Intracelluläre Pangenesis, a bottom-up determinist theory of organisms' development and function, as part of the strategic goal of his Die Entstehung der Arten durch Mutation. Indeed, the “unit character” was as much a notion of development as one of inheritance. As conceived by Bateson, unit characters were defined by properties that segregate according to Mendelian rules, and he tried to explicate their role in the physiology and development of the organisms accordingly (see Chapter 3).
Thus, from its inauguration the discipline of genetic research was burdened or blessed with the dialectic tension between the methodological reductionism of hybridists, deducing genes from traits, and the conceptual determinism of morphogenists, inducing traits from genes.
Although geneticists never lost interest in problems of physiology and development (Gilbert, 1998), they did adopt, as a rule, a bottom-up perspective of individual traits rather than a top-down perspective of organisms in their context. However, they needed some empirical handle to examine speculations such as the Weismann–de Vries hypothesis of specific nuclear particles being differentially exported to the cytoplasm of somatic cells.
Johanssen wished to discern the phenotype from the genotype (Johannsen, 1909). “Genes,” however, became what geneticists concluded Mendelian Faktoren had been. Were genes hypothetical constructs, autonomous structural entities or loci of differential functional emphases along an integral chromosome? Or, did all that not matter and genes were simply intervening variables, helpful entities for experimental work (see Chapter 4)?
Strictly speaking, there are no hypothesis-free concepts, and the distinctions between hypothetical constructs and intervening variables that proved very helpful historiographically, are more problematic when one attempts to apply them to personal conceptions. Whether we wish it or not, our concepts “are constructions in thought representing historically an immense amount of intellectual work … the scientific hypothesis does not come after the numerical data but before them” (Woodger, 1967, 366). Morgan, like Johannsen, insisted on presenting genes as intervening variables, with no hypothesis about the nature of the “something.” As late as in his Nobel talk he asserted:
Now that we locate [the genes] in the chromosomes are we justified in regarding them as material units; as chemical bodies of a higher order than molecules? Frankly, these are questions with which the working geneticist has not much concerned himself, except now and then to speculate as to the nature of postulated elements. There is no consensus of opinion amongst geneticists as to what genes are – whether they are real or purely fictitious – because at the level at which the genetic experiments lie, it does not make the slightest difference whether the gene is a hypothetical unit, or whether the gene is a material particle.
Concerning the matter of functioning of genes during development, I have contrasted … two possible views, and suggested a third. … It is known that the protoplasm of different parts of the egg is somewhat different, and that the differences become more conspicuous as the cleavage proceeds … From the protoplasm are derived the materials for the growth of the chromatin and for the substances manufactured by the genes. The initial differences in the protoplasmic regions may be supposed to affect the activity of the genes. The genes will then in turn affect the protoplasm, which will start a new series of reciprocal reactions. In this way we can picture to ourselves the gradual elaboration of differentiation of various regions of the embryo.
Morgan (1934a, 9–10)
In his classical paper of 1913 on the linear arrangement of sex-linked factors in Drosophila, Sturtevant described the phenotypes of the wings in combinations of genotypes: “The normal wing is RM. The rM wing is known as miniature, the Rm as rudimentary, and the rm as rudimentary-miniature” (Sturtevant, 1913b). The miniature phenotype is produced by a mutation in the rudimentary gene and vice versa, the rudimentary phenotype is produced by a mutation in the miniature gene.
This terminology, unlike the terminology we use today, had been introduced by Morgan: it reflected his sensitivity to the epistemological significance of the notation describing experimental results.
Similarity and variation among living creatures has long been a mystery that has stimulated the classification and organization of life into hierarchical systems, as well as the induction of the differential ranking of individuals in society. Variation and similarity also provided the material for one of the most basic ancient unfolding developments of human culture, animal and plant breeding. The first chapter of Genesis beautifully reflects the ancients' conception of the hierarchical, as well as anthropocentric, catalog of the universe:
In the beginning God created the heavens and the earth … Then God said, “Let the land produce vegetation: seed-bearing plants and trees on the land that bear fruit with seed in it, according to their various kinds.” And … the land produced vegetation: plants bearing seed according to their kinds and trees bearing fruit with seed in it according to their kinds. … And God said, “Let the land produce living creatures according to their kinds: livestock, creatures that move along the ground, and wild animals, each according to its kind.” … Then God said, “Let us make man in our image, in our likeness, and let them rule over the fish of the sea and the birds of the air, over the livestock, over all the earth, and over all the creatures that move along the ground.”
Genesis 1, 1–26
Animals and plants were believed to have been generated by God as distinct kinds at the time of Creation, and it was reproduction that maintained the link of similarity in living creatures.