Emergence of Modern Genetics in Spain and the Effects of the Spanish Civil War (1936–1939) on Its Development

Journal of the History of Biology 35: 111–148, (2002)

SUSANA PINAR
Dpto. Historia de la Ciencia
Instituto de Historia, CSIC
C/ Duque de Medinaceli 6
28014 Madrid
Spain

Abstract. The aim of this paper is to show how modern genetics reached Spain through the Junta para la Ampliación de Estudios e Investigaciones Científicas (JAE) during the decade of 1920s, the role played by key persons, and the level of development this discipline achieved from its different points of inception and under the conditions of financial scarcity and political turmoil that prevailed during the Spanish CivilWar (1936–1939). In addition, the effect of the war on the continuity of the lines of research already began is outlined, identifying the main area in which genetics first reappeared: agronomy.

Keywords: agronomy, genetics, JAE, Spanish Civil War, Spain

At the beginning of the 20th century, the absence of a research tradition together with a lack of commitment for its political and support prevailed in Spain. At that time, a promising period began with the establishment of the Junta para la Ampliación de Estudios e Investigaciones
Científicas (Board for Advanced Studies and Scientific Research, or JAE).

This institution, created in 1907 for the development of science, made the introduction of new disciplines easier and raised the scientific level of Spain through a policy of fellowships abroad. This pattern of science importation placed Spain among science’s peripheral countries. Thus, modern genetics reached Spain as early as 1921 through two JAE’s centers: the Museum of Natural Sciences of Madrid, where Antonio de Zulueta y Escolano’s laboratory was placed, and the Misión Biológica de Galicia (Galician Biological Mission) directed by Cruz Ángel Gallástegui Unamuno. A third center was established without much success in the Medical School of Madrid.

The JAE tried courageously to overcome Spain’s scientific stagnation and to fill the gap between Spain’s scientific activity and that of developed countries, but the incipient institutionalization of scientific research was interrupted by the outbreak of the Spanish Civil War (1936–1939) and the
SecondWorldWar. This interruption meant the disappearance of the previous research groups in genetics and the rebuilding of the discipline from different sources. Moreover, the golden era of classical genetics had passed, making way for molecular biology and molecular genetics.

The aim of this study is to analyze what made the introduction of classical genetics into Spain so difficult as compared with other European countries such as Great Britain, Italy, Scandinavia, Germany, and even USSR, where Mendelian genetics was successfully introduced up to the late 1930s and early 1940s. I will describe the political, social, and scientific factors that may have helped or prevented the introduction and institutionalization of genetics, focusing on the role played by key persons and under the conditions of financial scarcity and political turmoil that prevailed during and around the time of the Spanish Civil War. In addition, I will outline the effect that war period had on the continuity of the lines of research that had already begun and were institutionalized: agronomy. Accordingly, the paper will divided in two main parts, one dealing with the first genetics experiments accomplished on the field of biology, and the other with the application of genetics to the improvement of animals and plants.

Antecedents of the JAE and the Reception of Mendelism at Zulueta’s
Laboratory of Biology
The Spanish scientific structure of the first third of the 20th century was in many ways the outcome of the scientific institutions established at the end of the 19th century to overcome the power of the Church hindering the advance of science. The liberal revolution of 1868, which installed the short-lived First Republic in Spain, opened the doors to new ideas regarded as anathema by the
Catholic conservatives. At that time, Spanish society gradually became ideologically polarized and after the restoration of the Monarchy (1875–1923) Spain went into a constant alternation of conservative and liberal governments, in which university policy was determined along the changing political views. During that period, the increasing antagonism between liberal and conservative positions made it unwise to expound controversial ideas such as Darwinian evolution.(1) Consequently, as a result of the enactment in 1874 of a decree forbidding the teaching of anti-Catholic doctrines, many professors in favor of ideas such as Darwinism lost their university chairs. This was the case of the Krausist (2) philosopher and pedagogue Francisco Giner de los Ríos, who left the university along with other teachers and founded the Institución Libre de Enseñanza (Free Institute of Education) in 1876. The Institución taught the values of liberal democracy, social equality, and modern science, seeking to become the first non-state Spanish university and to reform all levels of education. However, the Institución failed due to lack of political
and social support. When in 1881 Giner de los Ríos and others were restored to their positions at the university, the Institución disappeared but not its reformist legacy, which would influence a new generation of politicians and intellectuals.

The Spanish-American war of 1898 had deeply affected the collective opinion of the Spaniard about themselves. It was not only the loss of the colonies (Cuba, Philippines, and Puerto Rico), but also the widespread feeling that the defeat was due to Spanish scientific and technical underdevelopment
as a result of the previous years of obscurantism. The positive side of this perception is that it created an environment propitious for the development of science, because the parties agreed to separate educational and scientific ideas from their ideological struggle.(3) The Regeneracionismo(4) movement emerged around the turn of the century, with the goal of solving the problem of stagnation at all levels. Four generations pushed to abandon the pessimism caused
by the defeat and to strive for the future. The generation of 1880, formed under the influence of the Institución, was the most involved in science development. The 1906 Nobel laureate, Santiago Ramón y Cajal, was its best known member and he became the first president of the JAE.(5)

The JAE was inspired by the previous Institución and was financed from the national budget as a public institution dependent upon the Ministry of Public Education. Trying to avoid the mistakes of the Institución, the JAE sought to stay away from the political fluctuations of the first third of the century, declaring itself an apolitical institution to promote education, opened to all ideologies. This allowed the JAE to outlive the dictatorship of Miguel Primo de Rivera (1923–1929) and benefit from the appearance of the Second Republic (1931–1939).

In response to the voices that sought to increase intellectual contact with scientifically developed countries, the JAE started a policy of granting fellowships to study abroad. This measure was followed by the reorganization and creation of scientific laboratories, mainly around Madrid, where grantees who returned home could carry on their research, introducing the new advances of
foreign science in Spain and invigorating their disciplines. Thus, the Laboratory of Biology of Antonio de Zulueta was established at the Museum of Natural Sciences in Madrid among a number of new laboratories created by the JAE and directed by outstanding scientists mainly trained abroad.

Despite the excellent work of JAE secretary José Castillejo, who supervised
the fellowship awards as well as the use of financial resources, scarcity
of funds was the norm in most laboratories.6 Consequently, international
support played and essential role in science development, as in the case of the
Rockefeller Foundation’s contribution to Spanish physics for the construction of the Instituto Nacional de Física y Química (National Institute of Physics and Chemistry).(7) The first references to Gregor Johann Mendel’s work turned up during the 1910s in the context of the apparent inheritance of some infectious diseases such as syphilis or tuberculosis, in relation to blood pathologies such as hemophilia, and in discussions about eugenics. As an example, in1911 José Chocomeli published an essay in a Valencian journal expounding his idea of inheritance after Mendel and Francis Galton, and treating both as complementary, rather than antagonistic theories. The same year, Antonio Lecha Marzo referred to Emil August Freiherr von Dungern’s work on blood groups, factors that he thought followed Mendel’s “laws;” Patricio Borobio Díaz wrote “La Heredo-Sífilis” (“The Hereditarian Syphilis”), and three years later the Academy of Medicine of Madrid organized a debate on the concepts of congenital and hereditary syphilis.(8)

Otherwise, at that time references to Mendel in Spain amounted to little more than a flirtation. At the end of the 1910s some Spanish cytologists began the first experimental studies on Mendelism. In countries where the Mendelian-Chromosome theory was introduced, its degree of acceptance has been related to the existence of a cytological tradition that could support the relationship between Mendelian factors and chromosomes, as well as to the presence of a scientist who had direct experience in Thomas Hunt Morgan’s laboratory.(9)

It is difficult to say that a cytological tradition existed in Spain prior to that moment. In fact, the JAE had no cytological laboratory, but opened several dedicated to histology that also did research on cellular biology. The Histological School created around the Nobel Laureate Ramón y Cajal proved his success in attracting researchers to biological studies. Members of this School had no active role in helping to introduce theMendelian-Chromosome theory, but their influence involved promoting histology and related sciences among Spanish researchers. In this, the most interesting scientists were a group of Jesuit biologists and the group that arose at the Museum of Natural Sciences in Madrid.

Father Jaime Pujiula i Dilme was the most remarkable member of the first group. After becoming a priest in 1906, Pujiula went to Germany and Vienna where he studied zoology and embryology. In 1908 he was entrusted with organizing the Ebro’s Biological Laboratory (Catalonia), later called
Sarriá’s Biological Institute when it moved to Barcelona. He was president of the Catalan Institution of Natural History (1925–1928) and member of the Biological Society of Catalonia. Pujiula was an antievolutionist and very critical of Mendelism. Between 1914 and 1919 he carried on breeding experiments on rats to verify Mendel’s works but could not repeat them, which was mainly due to Pujiula’s lack of control of the strains used in the crosses, which he ignored because he assumed that they were uniform (homozygosis) for the traits investigated.(10) At that time, he did not accept the existence of any kind of permanent or unchanging inherited biological units; instead he spoke
of some kind of centers modifiable by association, dissociation or chemical changes. Years later he would come to accept Mendel’s laws, the role played by chromosomes in sex determination, and Morgan’s theory, although with reservations.(11)

The second group was established at the Museum of Natural Sciences around Antonio de Zulueta y Escolano, who later became the central figure of Spanish genetics. Zulueta was born into a prosperous Barcelona family in 1885, and died in 1971. His two sisters were nuns and his older brother was Luis de Zulueta y Escolano, pedagogue and politician, who became Minister
of State of the Republican government of Manuel Azaña in December 1931.

In 1933 Luis de Zulueta was appointed ambassador to Germany, and three years later to the Vatican. Both brothers and Julián Besteiro, president of the UGT (General Union of Workers) and the PSOE (Spanish Socialist Workers Party), married Cebrián sisters. In early 1939, while Madrid was the only city still resisting Francisco Franco’s troops, Besteiro took part in the peace
negotiations representing the Junta de Defensa of Madrid. When the Spanish Civil War ended, he was imprisoned in Carmona (Seville), where he died as consequence of a bacterial infection for which he was refused medical assistence.(12)

After studying with the Jesuits in Barcelona, Antonio de Zulueta got a natural sciences bachelor degree, graduating in 1909 at the University of Madrid and one year later from the Sorbonne in Paris.(13) When Zulueta got his doctorate in November 1910, his career was directed towards the
zoology of protozoa.(14) A month after completing his doctoral dissertation, he received a one-year fellowship in Berlin to study reproduction of protozoa with Max Hartmann. However, in April 1911, he returned hastily before the end of his fellowship, when he was appointed interim curator of the
Museum of Natural Sciences by royal order of May 3, 1911.

At that time, this Museum was directed by the entomologist Ignacio Bolívar, also vicepresident of the JAE and the successor of Ramón y Cajal after his death in 1934. With Bolívar the Museum entered a new stage focused on conservation of collections and documents, research and spreading of knowledge by means of exhibitions and courses. In this context, the JAE entrusted Zulueta with the development of a biology course to train applicants for a fellowship abroad in the most common techniques of cytology and embryology. The training, offered uninterruptedly from 1914 to 1936, was structured in two sessions of 4 hours per week, sometimes complemented with lectures, lasting form October to April. Students from all fields of natural sciences attended the course, whose subject ranged from cytology and embryology to classic genetics.(15) Simultaneously, other institutions began to organize courses on inheritance taught by foreign scientists. As an example, in 1917 Leclerc du Sablon, from the University of Toulouse, taught a course on vegetal physiology at the Residencia de Estudiantes (Students Residence) in Madrid, including four lectures about “Hybridization and Heredity;” in 1919 l’Escola Superior d’Agricultura (High Agricultural College) in Barcelona invited Paul Dechambre, from the Veterinarian College of Alfort and the Agricultural
College of Grignon (France), to lecture on “Inheritance and its Application on Animal Production,” and in 1930 Julius Bauer taught about “Mendel Laws.”

As part of Zulueta’s course, a teaching “Laboratory of Biology” emerged, although without official standing until 1932 when Zulueta was appointed head of it by ministerial order of May 13. Among the first papers produced by the laboratory collaborators are those by Manuel Bordás Celma, visiting
fellow and director of the Escuelas Pías of Vilanova i Geltrú (Barcelona), on gametogenesis in the arrow worm Sagitta bipunctata and by José Fernández Nonídez on spermatogenesis in the beetle Blaps lusitanica.16 Both papers, published in 1914, dealt with the validation of the chromosome theory and, in particular, the second one with sex determination. Zulueta in those years was immersed in the verification of the cellular division patterns accepted for protozoa and the description of possible alternative explanations.(17) Later, between 1918 and 1919, Zulueta taught himself genetics, supposedly influenced by the direction cytological studies were taking on the problem of sex determination, in order to carry out his teaching duties properly. Almost
simultaneously, he began to develop basic experiments to confirm Mendel’s results in mice with different hair color patterns, experiments he repeated in 1925,18 and to breed the beetle Phytodecta variabilis. Products of this period are Zulueta’s translations of some evolutionary and genetic books:
The Theory of Evolution (1917) by William Berryman Scott, translated in 1920; On the Origin of Species (1859) by Charles Robert Darwin, in 1921, and A Critique of the Theory of Evolution (1916) by Thomas Hunt Morgan, in 1921.19 Afterwards, when Zulueta began his own research on Mendel’s
works, one of his first students, José Fernández Nonídez, got a fellowship to join Morgan’s group. Afterwarda, that stage Nonídez returned to Spain, and introduced the Mendelian-Chromosome theory.

Morgan’s Theory Reaches Spain. The Role of José Fernández Nonídez José Fernández Nonídez was born in 1892 in Madrid, but he spent most of his life as a teacher of anatomy at Cornell University in Ithaca, New York. He died in 1947, a few weeks after moving to the University of Georgia at Augusta. Nonídez achieved some international recognition for his histological work about the parafollicular cells, or C cells, of the mammalian thyroid and for his description of the glomus aorticum.(20) In Spain he is better known as the introducer of genetics, which began when Nonídez, then professor of zoology in the University of Murcia, applied for a fellowship to work at the Laboratory of Zoology and Comparative Anatomy of the University of Zurich.(21) The fellowship was granted by royal order on January 19,1917, but because of World War I.22 Nonídez requested that his grant be replaced by another to study cytology in relation to sex-determination and inheritance under the direction of Edmund Beecher Wilson and Thomas Hunt Morgan at Columbia University.(23)

In November 1917, Nonídez joined Wilson’s laboratory. There, he revised the preparations of the beetle Blaps lusitanica which he had brought from Spain, studying again its spermatogenesis and the X-complex.(24) He was able to trace the history of the chromosomes throughout spermatogenesis and solve the uncertainties remaining in his previous papers, where the
number of chromosomes and the composition of the X-chromosome were in conflict.(25) The illustration and study of meiosis in the spermatogenesis of Blaps were included in the 3rd edition of Wilson’s The Cell in Development and Heredity.(26) In addition, Nonídez learned the research methods used by Morgan for the study of inheritance; but, he did not participate in the gene
mapping project of the fruit fly Drosophila melanogaster. After renewing his fellowship for another year.(27) Nonídez assisted Wilson with his course on cytology, published an anatomical investigation of the reproductive system of Drosophila melanogaster, and collaborated with Charles William Metz as research associate in the cytological studies of the spermatogenesis of the fly
Asilus sericeus.(28) A previous collaborator of Morgan’s, Metz tried to create a fly group of his own at the genetics laboratory of Cold Spring Harbor in 1919, where he wanted to lure Alfred Henry Sturtevant and Calvin Blackman Bridges. Once the project was stopped, arrangements were made for the rival groups to cooperate and room was made in the Schermerhorn Hall for Metz,
Nonídez, Rebecca Lancefield, and a technician. In Sturtevant’s words “Not a poor group (. . .) but hardly the lot of first-class, highly trained geneticists that he [Metz] started out to get together.”(29)

When Nonídez returned to Spain in 1920, prepared to bring genetics back to that country, Zulueta and Ignacio Bolívar, carrying out the JAE’s project of scientific development, encouraged Nonídez to teach Morgan’s genetic ideas in a summer course. These lectures were published by the JAE
as Herencia Mendeliana (Mendelian Inheritance), the first Spanish text to include in detail not only Mendel’s works, but also the recent achievements of Morgan’s group: the Mendelian-Chromosome theory, linkage, crossingover, chromosome mapping, non-disjunction, etc. This book also included
two appendices with practical instructions for those who wanted to start experimental research on genetics of plants and animals, in particular Drosophila.

A couple of years later, Nonídez wrote Variación y Herencia (Variation and Inheritance), with great reception among agronomists and veterinarians from Spain and South America,(30) involving four editions between 1923 and 1946.(31) Meanwhile Herencia Mendeliana was published again in 1935 with lengthy modifications including the discoveries of the previous 12 years: the sex chromosomes of dioecious plants, intersexuality in the gypsy moth Lymantria and in Drosophila, epistatic and hypostatic phenomena, translocations, deletions, duplications and inversions of chromosome segments, etc.

When Nonídez went back to the United States the same year he had returned to Spain, looking for a better place to pursue his studies, Zulueta took upon himself the difficult task of developing genetics at the Museum of Natural Sciences. This effect, as we will see, brought new Spanish genetics
to an international scientific level.

Experimental Genetics at Zulueta’s Biological Laboratory
Zulueta’s greatest scientific result came after he began to investigate the inheritance of color types of the beetle Phytodecta variabilis.(32) The selectionof this insect affected the future of Zulueta’s career. By the 1920s, the fly Drosophila had displaced many other organisms from research with its short reproductive cycle (from 19 days to 3 weeks), producing up to four hundred offspring, and being cheap to feed. On the contrary, Phytodecta required a restrictive diet, living only on the fresh broom Retama sphoerocarpa, and producing one generation per year. However, Phytodecta had in its favor a distinguished morphological trait that seemed to be linked to sex: the different color types of the outer hard wing-case of beetles or elytra. With the beetle Phytodecta Zulueta got the first evidence of the existence of a non-recessive character linked to a Y-chromosome cytomorphologically different from the rest of the chromosomes or autosomes.(33) This discovery was the reason why Zulueta never abandoned this beetle in spite of the meagre scientific results.
Zulueta’s laboratory kept some Drosophila mutants for teaching and student practice, but for unknown reasons he did not use this organism for research. This kept Zulueta’s laboratory separate from the global community created around Drosophila.

As regards Zulueta’s research, Wilson’s studies on the role of chromosomes in sex determination defined two main arrangements of sex chromosomes. In the first type, females had a pair of equal sex-chromosomes called X (homogametic sex, XX), while the male lacked one of the members of this pair (heterogametic sex, XO) or had a second, smaller chromosome, called the Y, paired with the X (heterogametic sex, XY). Because all degrees of X and Y-chromosome combinations could be found among different species – from the equality of both chromosomes to the complete absence of the Y –, the general opinion was that the Y-chromosome contained no significant genetic elements. Morgan stated that those observations did not show the Ychromosome to be empty, except in the sense that it contained no genes that behaved as dominants to recessive genes of the X-chromosome.(34)

Because the first observations furnishing genetic evidence that the Ychromosome could carry dominant genes and undergo crossing over were made in fish, where the Y-chromosomes are cytologically and morphologically undistinguishable from the autosomes, Morgan rejected those cases studied by Johs Schmidt (1920) in the cyprinodontid fish Lebistes reticulatus, by Tatuo Aida (1921) working with the fish Aplocheilus, and by Öjvind Winge (1922, 1923) also in Lebistes. In all these fishes the females of different races were closely similar in coloration while the males were differently marked. When the male of one race was bred to a female of another race, the sons were colored like the father. If these hybrids were inbred, the sons were again like the father. The color was not inherited from the mother, and the same happened in later generations. In these cases, where sex-chromosomes were indistinguishable from the autosomes, Morgan argued that another
possible explanation was to assume that: “The sex-chromosome may be a part of, or have become attached to, a pair of autosomes. This means no more than that one part only of the ‘sexchromosomes’ has the function of an X- or Y-chromosome, which means again that the Y-part is different from the X-part, and that no crossing-over takes place between these parts (. . .) this mechanism gives the same kind of result, so far as sex is concerned, as does the occurrence of independent sex-chromosomes.”(35)

In the case of Zulueta’s beetle, the Y-chromosome is morphologically distinguishable from the autosomes.36 The heredity of the four color types of Phytodecta37 was explained by Zulueta under the assumption that the chromosomes that carried the genes were the sex-chromosomes and that the genes could be present in the X’s, and any one in the Y’s. Against this, Morgan argued:
If these genes are in the sex components which, ex hypothesis, do not interchange, the question arises as to how the same kind of allelomorphs may be present in the X- as well as in the Y-component. One answer would be that they have arisen from the same kind of original gene as
independent mutations, sometimes in the X-component, sometimes in the Y-component (. . .), but another explanation will do as well. Suppose, for instance, that the genes for the four color markings are not present in the X- and Y-components but in the autosomal components and that they are
allelomorphs. The results will then be explicable on the assumption that there is either no crossing-over in the male or that it is so infrequent as not to have arisen in Zulueta’s [experiments].(38)

Now we know that Zulueta was right. The heredity of the color types of Phytodecta was a case of partial linkage to sex, in which the locus of elytrum color was placed on the homologous segment of the X and Y-chromosomes. An analysis of Zulueta’s study establishes that on the crosses with black
heterozygotic males XBYY – B and Y are alleles for color black and yellow –, the beetles only produced gametes XB and YY, when theoretically they could also produce gametes XY and YB, that is, those resulting from a crossing over. This means that the locus implicated was close to the place where the chromosomal segments became different in both chromosomes, so the
crossing-over rate was almost zero.(39)

One year later, in 1926, Curt Stern provided further evidence for Zulueta’s results about the existence of dominant genes in the Y-chromosome, working on Drosophila. At that time (1924–1926) Stern was at Columbia University with Morgan, supported by the Rockefeller Foundation. His first Drosophila problem was to study the inheritance and sexual expression of the mutant recessive gene “bobbed” that produced shortened bristles on female flies, but not on males. This peculiarity was solved by finding that some females had 2X chromosomes as a result of non-disjunction, but had been fertilized by a sperm bearing a Y-chromosome, making them XXY; in these females the
bobbed character was suppressed as in males. Clearly, the Y-chromosome, normally found only in males, had to carry normal allele of the bobbed mutant that suppressed its expression.40 These results were an irrefutable proof that the Y-chromosome was not entirely lacking dominant genes. The research on the effects of fragmentation and translocation of parts of the Y-chromosome
would be completed only after Stern had returned in 1926 to Richard Benedict
Goldschmidt’s laboratory in Berlin-Dahlem, where he also demonstrated the
cytological basis of crossing over.

Zulueta’s was the first original study in genetics from Spain to reach the international community. Not only did Morgan cite Zulueta’s discovery in the 1926 paper mentioned and in The Scientific Basis of Evolution,(41) but in the following years other authors such as Wilhelm Johannsen, Emile
Guyénot, Curt Stern, and Björn Föyn, reported Zulueta’s data, illustrations, and genealogies when dealing with heredity linked to the Y-chromosome or with multiple allelomorphs.42 A few years later, Zulueta would undergo another “immersion” in Morgan’s genetics by means of a Del Amo Foundation(43) invitation to go to California for lecturing and further training.(44)

Thinking that it could be beneficial for his teaching at the Museum, Zulueta decided to go to the California Institute of Technology, where Morgan’s group had moved in 1928.45 For personal reasons, Zulueta postponed his trip until February 1930 and remained in Pasadena through April.46 There, he learned, under the supervision of Bridges, the techniques of chromosome mapping
designed by the group in 1911 and conducted his only investigation with Drosophila, locating with greater accuracy the locus “light,” one of several involved in the determination of the red color of Drosophila’s eyes. Zulueta’s work at CalTech manifests his efforts to stay in touch with mainstream
genetics. In Spain, Zulueta’s and Bridges’ papers about the localization of the mutant “pink-wing” of Drosophila, were published in the same number of the entomological journal Eos.47 On the American side, the Carnegie Year Book of 1929–1930 reviewed Zulueta’s achievement.48 Two years later, Zulueta returned to the United States on the occasion of the VI Congress of Genetics in Ithaca, where he showed some samples and drawings of Phytodecta, illustrating the different color patterns transmitted through the X and Y-chromosomes.(49)

As with many others scientific fields, genetics was strenghthened with the arrival of the Second Republic in 1931. One year later, under the sponsorship of a private institution, the Conde de Cartagena Foundation that operated in the Real Academia de Ciencias, Exactas, Físicas y Naturales (Royal Academy of Mathematics, Physical, and Natural Sciences), the first chair of genetics was created and given to Zulueta.(50) Previously, by royal decree of August 25, 1926, the reform of the studies of Bachillerato (more or less the equivalent to high school) took place with the direct mediation of the Museum of Natural Sciences, succeeding in including the study of basic
genetics within the subject of biology. A year later, genetics appeared in high school textbooks(51) and, later, by royal decree of March 19, 1928, genetics entered the Natural Sciences Schools as part of the biology degree program at the University, but no chairs in genetics were created. The situation was similar at the Agronomic and Veterinarian Colleges, but genetics entered their degree program four years earlier, in 1924. The first course of the Conde de Cartagena chair was taught at the Museum, later moved to the Academy and, from 1934 onward, it was taught at the University in Madrid.(52)

After 1934 there are no publications of Zulueta until the end of theSpanis h CivilWar (1939), get neither he, nor the laboratory was inactive. The study of Phytodecta continued, while Zulueta’s student Fernando Galán(53) began to breed different varieties of the four-o’clock Mirabilis jalapa and
the snapdragon Antirrhinum majus and, later, different varieties of the “explosive” cucumber Ecballium elaterium.(54) In addition, Zulueta prepared a detailed report “Die Spanischen Eselrassen” (“About the Races of Spanish Donkeys”), at the request of Emil Fedotovich Liskum from the Institute of Animal Husbandry of Moscow.55 The scant publication record was due to the difficulty of Phytodecta research, the searching for new organisms of investigation, the huge teaching duties for meagre salaries, and the Civil War. Besides his very important teaching assignment, Zulueta also took part in the popularization of genetics, carrying out collaborations in Revista de
Pedagogía and Conferencias y Reseñas Científicas56 and writing a large number of scientific reviews mainly for the Boletín de la Real Sociedad Española de Historia Natural. In addition, from 1927 to 1936, he was chief of the scientific journal Investigación y Progreso, where he translated around thirty papers on genetics from German.57 He took charge of the new publication
Revista Española de Biología, and also was the correspondent in Spain of the German journal Berichte ueber die Wissenschaftliche Biologie and the Dutch Resumptio Genetica. For his outstanding collaboration in the improvement of Spanish and German scientific relationships, Zulueta was awarded the Merit Cross by the German Red Cross during Hitler’s regime.(58)

Contributions of Genetics to Human Inheritance and Medicine
Zulueta’s translations of Biology of Twins (1917) by Horatorio Hackett Newmann in 1922 and Educational Biology (1930) by William Lewis Eikenberry and Ralph Augustus Waldron in 1931 suggest that he was interested in some aspects of “eugenics.”(59) But, what was the relation of Spanish genetics with eugenics? As in France and in many Latin American countries, in Spain
eugenics took a more hygienic than hereditarian turn, being more concerned with health reform than on the promotion of the sterilization of criminals or the feebleminded. Those measures were only supported by some Spaniards who had an early contact with the United States, such as Nonídez or the psychiatrist Gonzalo Rodríguez Lafora.(60)

Zulueta was always very cautious on these matters and never spoke in favor of any eugenic methods, although he showed some interest in the most genetic aspects of eugenics. In addition to his many translations, around 1935 Zulueta began to study some cases of pathological inheritance in
humans, such as lobster claw deformity (ectrodactylism), extra fingers or toes (polydactylism), and some skin problem of the hands and feet (hyperkeratosis). (61)

Later he went as an official delegate to the III International Congress of Eugenic celebrated in New York in 1932, but made no contribution. However, one year earlier, he had an active participation on the I Jornadas Eugénicas Españolas (1st Spanish Eugenic Conference). Among the program
of this conference two practical courses on genetics were entrusted to Zulueta and the physician Jimena Fernández de la Vega. Zulueta’s lecture entitled “Herencia en Animales y Plantas” (“Heredity in Animals and Plants”), dealt entirely with genetic topics: Mendel’s laws, pleiotropy, multiple allelomorphism, and the effects of inbreeding and crossbreeding in relation to some human pathologies, while Fernández de la Vega focused on human heredity, dealing with Mendelian transmission of psychological characters, where she repeated some clichés about the genealogy of genius and feebleminded, and on the significance of mutations as factors of evolutionary progress and sources of diseases.(62)

In fact, Fernández de la Vega made the only serious attempt to take genetics into the field of medicine, where the concepts of inheritance were linked to the sense of predisposition or what was referred to as constitutional types. Fernández de la Vega was the disciple of the parasitologist Gustavo Pittaluga and physician Roberto Nóvoa Santos.63 With the support of the JAE, she was trained in Berlin where she studied biometry with Friedrich Kraus and Thedor Brugsh, in 1924–1925. Later, she spent one year in Hamburg with Hermann Poll, where she verified Morgan’s experiments of sex-linked inheritance in the white-eyed Drosophila and carried out a study about pathology
and inheritance of identical twins.64 Then, she stayed six months with Julius Bauer in Vienna, where she studied constitutional types, and later, between 1933–1934, attended the school of Nicola Pende in Genoa. When Fernández de la Vega returned to Spain in 1927, she began
experimental research about inheritance of blood groups and several blood pathologies, where genetics ideas were better understood.65 In 1930 she translated Julius Bauer’s book Konstitutions und Vererbungdslehre (Constitution and Heredity), from its second edition of 1923, and after her association with Nóvoa Santos, she was nominated for the direction of the Sección de
Genética y Constitución (Section of Genetics and Constitution), which would be created in 1933 in Madrid, as part of the pathology chair of the School of Medicine, for Nóvoa Santos. When Nóvoa Santos died that same year, the Sección was reduced to a theoretical seminar. Later, Fernández de la Vega also taught a genetics course (1934–1935) at the Gregorio Marañón’s chair of the Instituto de Patología Médica (Institute of Medical Pathology), whose lectures were published as a book titled Herencia Patológica en la Especie Humana (Pathological Inheritance in the Human Species), becoming the first Spanish text about human genetics written by a physician. Despite this, genetics did not take root in the School of Medicine. Although genetics was included in the medicine degree program within the subject of “citology, embriology and genetics” during 1960s, nowadays there is not a genetic chair yet.

Impact of the Spanish Civil War on Zulueta’s Genetic Research
Back to Zulueta and his research: just before the Spanish Civil War, genetics had acquired its place among the subjects of growing interest in Spain, although equipment was scarce and space was limited. The situation could have changed drastically, because the Rockefeller Foundation turned its eyes to Zulueta’s Laboratory of Biology. The Foundation agreed to contribute to equipping a new laboratory of genetics, although not to the construction of a new one,(66) while the JAE had to assume the purchase of specialized journals, as well as paying for an assistant to Zulueta as head of the laboratory.(67)

The new assistant was going to be Käte Pariser, an early student of Richard Goldschmidt, who had been in Zulueta’s laboratory for three years, studying the origin of the malformation of gonads and the pelvic extremities of the newt genus Triturus and their possible association with the almost total lack of males in some interspecific hybridizations.68 When the social situation in Spain deteriorated preceding the outbreak of the CivilWar, she left Spain and went first to Tel Aviv and later to Sydney. In addition, Fernando Galán got a fellowship abroad from the Rockefeller Foundation, planning to go to Caltech with Morgan to carry out research on the application of the silver staining method of the Spanish School of Ramón y Cajal to the giant chromosomes of the salivary glands Drosophila.(69) These huge chromosomes, discovered by Theophilus Shickel Painter, were excellent material for the observation of phenomena such as duplication, translocation, deletion, and inversion of
chromosomal segments. Eventually, Galán declined the fellowship because he thought his duty was not to abandon his country at war, and because he wanted to continue his research collaboration with Zulueta.

The financial aid that the Rockefeller Foundation offered to Spanish genetics could have been crucial because, in countries such as Brazil, it insisted that the government participated in the creation of full-time positions so that scientists be sufficiently well paid and would not need to have
additional jobs in order to earn a living;70 a problem that affected to Spanish researchers at least until the late 1950s. Moreover, the Rockefeller Foundation always gave financial support, facilitated the exchange of researchers, and, as in the case of Brazilian genetics, promoted a international team research, which yielded better results than those obtained from the independent and
uncoordinated efforts of several researchers working in isolation.

A little before the beginning of the Spanish Civil War in 1936, Zulueta and Galán started a genetic investigation of the “explosive” cucumber Ecballium elaterium. This plant of the gourd family exhibited monoecious and dioecious subespecies, fertile when hybridized with each other or with
their hybrids. Several young scientists were participating in the laboratory’s work, observing and recording the experiments of crossbreeding between subspecies and hybrids of Ecballium, using as their experiment field what in other times had been the athletic fields of the Students Residence.(71) After the war, this research was continued by Galán at the University of Salamanca, where he carried on “the first complete genetic and experimental analysis of the zygotic monoecia and dioecia,”(72) after the theoretical method that Carl Correns had proposed in 1907 for Bryonia dioica, another species of the gourd family.

Among the efforts to give continuity to scientific work during the Civil War (1936–1939), Zulueta participated as Spanish delegate to the Congrés du Palais de la Découverte celebrated in Paris in 1937. He took advantage of being abroad to write to Gregorio del Amo in order to reiterate his gratitude for the opportunity he gave him to go to California in 1930.(73) Del Amo’s answer was to offer him a new opportunity to come back to CalTech and get acquainted with the genetic advances of the last six years.(74) Neither at this time, nor later, it seems that Zulueta thought about moving abroad, although many other Spanish scientists, as well as his own brother Luis, left as a
consequence of the Civil War. Thus, during the Civil War and after the exile of Bolívar, Zulueta remained in charge of the Museum of Natural Sciences, becoming its interim director.(75) The situation of the laboratory during the Spanish Civil War was well reported by John Burdon Sanderson Haldane, who wrote in Nature: "During a recent visit to Madrid (. . .) I was able to visit some of my
collegues there, and was delighted to find that research work in genetics was continuing. Prof. A. de Zulueta, in the intervals of hiding the more precious contents of the biological museum in cellars, was continuing his work on the polymorphic beetle Phytodecta variabilis. Prof. Galán, of Salamanca, was very appropriately breeding the “explosive cucumber,” Ecballium elaterium. (. . .) Our discussion of these topics was interrupted by an air raid considerably more severe than any of those on London in
1914–1918. However, no bombs fell very near us, and at the time I left, the Museo de Ciencias had not yet shared the fate of the University city, the Prado and the Museum of Anthropology. I think that the persistence of Zulueta and Galán under conditions which are, to say the least, uncongenial
for research, deserves to be recorded, and augurs well for the future of biology in Spain.(76)

However, Zulueta’s research did not count in Spain with the support of many followers, perhaps because they did not see an immediate practical application for it. This reason, in conjunction with the fact that the new genetic techniques, such as those for the construction of pure lines and
hybrids were already familiar to the investigators dedicated to the improvement of animals and plants,(77) explains why genetics found its most congenial home in the field of agronomy.

Genetics and Selection of Animals and Plants
At the beginning of this paper, it was said that at the turn of the 20th century Spanish science was stagnated, a phenomenon that reached other sectors of Spanish society. Thus, agrarian Spain was not very productive and ownership of land was characterized by extremes of size and provincial distribution. In some regions, such as the North, small holdings prevailed and were generally
insufficient to sustain peasant families, while, to the South, large holdings or latifundios redominated. Agrarian reform was a necessity, but due to its great difficulty it was postponed each time and it was badly done when it was eventually undertaken. Thus, in rural areas where large owners monopolized the land, unrest was endemic and this continuous agitation upset political life throughout the whole nation.(78) Industrialization had come late and was limited, but created a huge working class of low economic level. Many social organizations and trade unions, primarily of Anarchist and Marxist ideologies, seeking better conditions for workers, elicited a severe rebuff from various sectors of society. The government did not approach the question with intelligence and energy, which increased resentment and unrest during the first third of the century, eventually leading to the Civil War. Another area of conflict was the regionalism of Catalonia, the Basque Country and,
to a lesser degree, Galicia, which turned into nationalism, sometimes with an open separatist trend that seemed a menace for the nation.

Nevertheless, the underdevelopment of Spanish agriculture could have been used as a prime reason for the development of science and as a positive influence for the introduction of genetics. In other parts of the world, such as the United States, the rediscovery of Mendel’s works in 1900 brought new hopes for agronomy, so that what had been the “art” of the selection of animal and plants, sought to become a science. However, the revolution would not come for another twenty years or so, until the discovery, independently by Nils Herman Nilsson-Ehle and EdwardMurray East, of the phenomenon now known as multiple factors, because it provided a proper interpretation within
the Mendelian framework for quantitative or “blending” inheritance, of great importance to breeders and farmers. The Darwinian concern for selecting better-adapted individuals was brought together with the Mendelian analysis of hereditary differences, following the ideas developed by Morgan’s group.

Hybridization came increasingly to be used in conjunction with selection.(79) The most important achievement of that time was the new technique of double hybridized corn, which played a vital role to overcome the Great Depression of the 1930s in the United States. Double hybrid corn was
introduced into Spain during the decade of 1920s. It could have helped to overcome some problems of production of Spanish agriculture, but its development was limited. The pattern of introduction of double hybrid corn technique and that of Morgan’s theory were both imported through the JAE
policy of fellowship abroad. Cruz Ángel Gallástegui Unamuno played the role of introducer for the hybrid corn.(80) Gallástegui, born into a family of Basque farmers in 1891 and died in
1960. He was trained in France (1909–1911) and Germany (1911–1914),where he got a degree in Agronomy from the Royal Agricultural College of Hohenheim-Stuttgart (Württemburg) in August 1914. In Germany he met Juan López Suárez, an influential physician who later married the sister of the JAE’s secretary José Castillejo, who determined Gallástegui’s future career.
In November 1917, López Suárez convinced Gallástegui to visit Morgan together, who advised them to go to the Bussey Institution at Harvard University, where Edward East(81) rapidly involved Gallástegui in corn research. It was at this point that the JAE awarded Gallástegui a fellowship.

It is known that hybrid corn was the combined effort of many scientists.(82) The idea of crossing inbred strains of corn was first proposed by George Harrison Shull in 1909,83 when he identified some important factors that became standard operating procedures in the corn breeding programs initiated later. East and Donald Forsha Jones worked on the effects of inbreeding and crossbreeding at the Connecticut Agricultural Experiment Station in New Haven, where in 1917 Jones carried out the experiment of crossing Shull’s simple hybrids. Five months after Gallástegui joined Harvard University in December 1917, East sent him to the Connecticut Station in order to help Jones in his research. The double hybrid seed was sown during the spring of 1918 and by harvest time they witnessed the first double hybrid corn of great vigor. After that, Gallástegui and Jones published together one paper analyzing some linked characters in corn and another paper popularizing the double hybrid techniques in the United States. Simultaneously, Gallástegui
published for the first time those results in the journal El Cultivador Moderno of Barcelona.(84)

Hybrid corn technology was adopted in other areas of the World quite rapidly following World War II; although it had reached Spain at the beginning of the 1920s, after Gallástegui returned in 1921. At that time, his old friend López Suárez was negotiating with the Real Sociedad de Amigos de Santiago de Compostela (Royal Society of Friends of Santiago de Compostela) to request from the JAE the creation of a research center dedicated to the genetic improvement of animals and plants in Galicia. This proposal, in accordance with the JAE guidelines, had been designed to
decentralize research, creating new laboratories in geographical areas that were able to help their establishment with financial aid.85 In April, 1921, the Misión Biológica of Galicia began its life under the direction of Gallástegui.

Its aims were the application of genetic techniques to the improvement of plants and animals, with a few doses of genetic training. The Misión focused on a few lines of research with direct application to the economy and agriculture of Galicia, such as corn research. Besides, the Misión also managed
to solve one of the biggest problems of seed experiment stations, namely the scarcity of land and the distribution of seeds, with the establishment of a cooperative. After years of experimentation with hundreds of inbred lines from Spain, America, France, and Italy, the Misión achieved three
double hybrids acclimated to Galicia’s different climates. These were the first achieved in Europe as early as 1927, and commercialized in 1930. They reported increments in production of around 5,000 kg/acre from the previous 3,500 kg/acre, reaching sometimes increments of nearly 20,000 kg/acre when combined with fertilized soil.(86)

The institution passed through two difficult periods, largely coinciding with those in political life: the dictatorship of Miguel Primo de Rivera and the Second Republic. Prior to and during Primo de Rivera’s government, stockbreeding issues were part of the Dirección General de Agricultura
(General Directorate of Agriculture). Some attempts to improve animal sector were made by two royal decrees in 1924, giving responsibilities to the Estaciones Pecuarias (Cattle Stations), reorganizing the Instituto Agrícola Alfonso XII (Alfonso XII’s Agricultural Institute), and leaving stockbreeding in the hands of agronomists, a very corporative collective, ready to fight for
its prerogatives and jurisdictions. That same year, genetics was included into the curriculum of the Instituto Agrícola Alfonso XII.87 No special chair was created but its teaching was given to Genaro Alas. Alas frequented Zulueta’s laboratory, where in 1927 he began a research on Drosophila.88
Also in the agronomic field, the Instituto Nacional de Investigaciones y Experiencias Agronómicas y Forestales (National Institute for Agronomic and Forest Research and Experimentation) was established in 1926 to coordinate the research stations scattered throughout Spain. Genetics was
incorporated in two main and important centers. The first one was the Estación Central de Ensayo de Semillas (Central Experimental Station of Seed) founded in 1907 for the improvement of wheat. In 1926 Antonio Esteban de Faura began the station’s modernization, applying new techniques
on inbreed lines from the Swedish Station of Svälof (Sweden) to wheat selection.

The second one was the Instituto de Cerealicultura (Cereals Institute) established in 1929 and directed by Marcelino Arana. This center counted with a laboratory and 9 experimental fields in different parts of Spain, and with Ramón Blanco and Alonso Ruíz de Arcauete among its staff. Around 1930 the Instituto paid the Misión for the genetic training of two agronomists:
Vicente Boceta and Miguel Odriozola, who later completed their training abroad. Boceta studied genetics with Erwin Baur in Germany and after his return continued working in corn selection.(89)

Odriozola went first to the Agricultural College at Cambridge, later to the Animal Genetics Institute of
Edinburgh and finally to the Rowett Research Institute in Aberdeen, returning to the Misión in 1933.
These centers experienced difficulties in supplying seed to the whole nation. A solution repeatedly attempted was the creation of cooperatives, but none except the Misión carried out the project. In 1930 Gallástegui and Daniel de la Sota created a Sindicato de Productores de Semillas (Trade
Union for Seed Producers, SDS), which was the first European institution to produce and sell hybrid corn seed. In addition, for Gallástegui the SDS was the best way to change Galicia’s old agrarian structure and transfer laboratory results into the field, introducing hybrid seed throughout Galicia and Spain.(90)

In previous years, the Misión had given for free, or for a very low price, double hybrid seed to the farmers, but its limited fields were not enough to provide the increasing request for seed.91 Thus, land from all its members was used, while foremen trained by the Misión supervised cultivation procedures for quality control of seed.92 With the SDS, the Misión became much more
independent from government subventions and political fluctuations, which had diminished in previous years. In 1933 a bulletin of the SDS was created to keep members informed of agricultural novelties. Three years later, the SDS, which had 216 members, mainly agricultural societies, expanded its management to other products and sold seed all over Spain.(93)

With the arrival of the Second Republic in 1931 a new period opened. The agrarian reform went into the foreground of political activities, where the problem of production joined the serious problem of land tenure, and stockbreeding also played a role in the debates. The Republican period
(1931–1939) meant a great impulse in all sectors related to genetics. As it was mentioned, the Misión achieved its peak after the creation of the SDS, expanding its investigation to other products such as potato, rye, and the selection of cattle and pigs. The Misión bought a herd of Large-White pigs as part of a project of porcine improvement for Galicia, carried out by Odriozola
after his return to the Misión in 1933. Studies of consanguinity and nutrition were made on this herd and they have been used as stud for breeders all over Spain. Nowadays, this herd is considered the second oldest of its kind in the world.(94)

The politicization of ideas and institutions during that period was important. At a regional level, due to the favorable attitude of the Republic towards the diversity of the Spanish regions, the Galician regionalists unfolded a tireless activity to achieve autonomy. The Partido Galegista (Galician Party, PG) included in its political program Gallátegui’s economic reform, based on the increase of production using hybrid corn and fertilizers. This initiated a series of reforms at all levels ending in the promotion of milk cattle to create a potent industry of dairy products to move part of the rural
populations to the cities.95 The PG’s adoption of the Misión as its symbol gave soundness and scientific respectability to its proposition. Moreover, the good results of the Misión and the SDS provided a successful propaganda, reinforcing Galicia’ self-confidence as an independent entity and providing a proof of the viability of its self-government.(96)

At a national level, the socialist deputy Félix Gordón Ordás(97) was initiating the reform of livestock breeding in 1931. This was a problem closely related to the establishment of veterinarians as a professional class, in which it was not only necessary to renew their knowledge, including genetics, but also to update their role in society, previously limited to sanitary matters. In 1931, Gordón inspired the creation of the Dirección General de Ganadería (General Directory of Animal Husbandry, DGG) after the model of the North American Bureau of Animal Industry.(98) The DGG unified in a single institution the administration and all the related centers, removing its responsibilities from the agronomists. This body was divided in three sections: one dedicated
to animal hygiene and health, the second to improvement and research, and a third to education and social work. The section of Fomento Pecuario (Animal Improvement) was until the war the responsibility of two General Inspectors: Gallástegui and Juan Rof Codina, also founder member of the Misión. Although Gallástegui got the degree of veterinarian around 1923, he had never practiced it, a condition needed to get the inspector post. Gordón wanted so much to incorporate Gallástegui to his team that a decree was issued to overcome the obstacle. Due to the rebuff of the veterinarian
community, Gallástegui left the post in January 1932, but the balance of his period was positive and his legacy was a normative for stud breeding. In May 1932, Gallástegui entered the Consejo Ordenador de la Economía Nacional (Coordinating Council of the National Economy) formed by fifteen members with advisory functions to the Ministers Council.(99) In 1933 the legislation
for stud books and milk production was modified, but the results were limited for irregularities in data register, prolonging the subjective method of morphological evaluation.

In education, Gordón proposed a complete reform of veterinary program in 1933. This included genetic training as part of the university and doctorate curriculum, but when in January 1932 the veterinary colleges reverted to the Ministry of Public Education, the project failed. However, genetics continued being taught as part of other subjects at Madrid by Juan Homedes Ranquini,
Fernando Hernández Gil, and José Ocáriz Gómez, at Barcelona by Pere Martir Rossell i Vilar, in Cordoba by Gumersindo Aparicio Sánchez and in Zaragoza by Rafael González. No other text books than Nonídez’s were written about that time, but in 1934 Gregorio Ferreras translated for the
journal La Nueva Zootecnia the book Animal Genetics (1925) by Francis Albert Eley Crew.

The veterinarian group teaching in Madrid was also part of the Instituto de Biología Animal (Institute of Animal Biology) established in 1931 as part of the DGG. Three great lines of research were established in the Instituto:Animal physiology and “zootechnology” (animal improvement), animal
pathology, and what it was called contrastación, dedicated to help breeders in the selection of fodder and quality inspections. The first section was directed by Homedes Ranquini and was focused mainly on genetic and cytological aspects of reproduction, as well as on animal endocrinology. Towards 1933– 1934, the veterinarian Leopoldo Calvo Sánchez, who had been training with Zulueta on a fellowship of the Ministry of Agriculture, began research on Drosophila in the Instituto.100 In 1933 the Instituto also began to publish its own journal, Trabajos del Instituto de Biología Animal. A few years before, in 1929, another specialized journal, La Nueva Zootecnia appeared, where a good number of papers on cytogenetics and genetics were published.
These journals collected information about courses and conferences on these matters, organized by different institutions such as the Veterinarian Colleges of Vizcaya and Zaragoza, the DGG or the Turró’s Association, being taught by Rafael González, José Ocáriz, Santiago Tapias, Isidro García, Homedes Ranquini, Zulueta, etc. Simultaneously with the veterinarian reforms, agronomy
was reorganized with the establishment in 1932 of the Instituto de Investigaciones Agronómicas (Institute of Agronomic Research, IIA). Its initial nucleus consisted of the Instituto de Cerealicultura, mentioned before, and the different stations dedicated to plant pathology, seed production, etc.,
which counted 64 centers in all, where genetics took a place to a larger or lesser degree. Its aim was to coordinate research centers and the numerous projects at hand. The IIA continued after the war, but it was reorganized and “National” added to its name, becoming the INIA. In 1935, it began to publish the Boletín del Instituto de Investigaciones Agronómicas.(101)

War and Postwar
The Civil War damaged deeply the incipient Spanish scientific structure that had begun to rise at the beginning of the 20th century. Many scientists died or went into exile, while those remaining were purged after the war to clear their political affiliations by law of February 10, 1939. Zulueta was accused of being affiliated to the Izquierda Republicana (Left Republican) party and loyal to its leaders. He was described as a friend of Ignacio Bolívar and as director of the Museum of Natural Science was charged with signing a manifesto in favor of the Republican president Manuel Azaña, and another against the bombardment of Madrid. In addition he was accused of attending the Paris Congress and returning afterwards to the republican zone with his family.(102)

Zulueta was found guilty and forbidden to enter his laboratory. A year later, he was allowed to work there again, but did not regain his salary until much later. Galán was also purged, but allowed to continue as professor at the University of Salamanca, where he moved the Ecballium plantations.
Gallástegui and Fernández de la Vega were accused but their cases were dismissed. Many others had worse luck. In Zulueta’s case, the severity of his sentence was, without doubt, due to his familial relationships (Luis de Zulueta and Besteiro), and his closeness to Ignacio Bolívar and the JAE.
The JAE disappeared, but its institutes became part of the Consejo Superior de Investigaciones Científicas (Spanish Research Council, CSIC), the new organization for fostering, directing, and coordinating scientific research, established by law of the 24th November of 1939. As the JAE
did, the CSIC maintained its independence from the university, which was in a very sensitive situation due to a serious drain of faculty, who had left the country because of the war. During the years following 1939, the CSIC counted with more resources, support, and staff than the university.
However, centers close to the central figures of the JAE, the Ramón y Cajal group, and the Museum directed by Ignacio Bolívar, were chastised by the military government. The Museum of Natural Sciences was abandoned for many years: Zulueta’s laboratory kept only a nominal existence. Later, the Spanish Academy of Sciences through its secretary José María Torroja Miret requested Zulueta to resume his teachings at the genetics chair of the Conde de Cartagena Foundation, where he remained from 1946 to 1952.(103)

During the rest of his active life, Zulueta would make only a few scientific contributions, playing an honorary role in Spanish genetics.104 Zulueta had numerous students, but no school of geneticists grew around him. Some of his students remained in the country and contributed to the development of genetics in Spain, most notably Fernando Galán. In Salamanca Galán facilitated Francisco J. Ayala’s emerging interest in genetics. Galán introduced Ayala to Zulueta around 1960, who in turn introduced him to Theodosius Dobzhansky, whom Zulueta had met during his stay in CalTech. Ayala remained in the United States, with successive faculty positions, but in close contact until
their deaths with Zulueta (1971) and Galán (1999) and, in a way, he is the strongest link to the early period of genetics. Eugenio Ortíz, who also was Zulueta’s student, became his successor at the laboratory of biology of the Museum of Natural Science.

Gallástegui continued in the Misión Biológica of Galicia until his death in 1960. Corn research was also done in the Cereal Experiment Center of the now Instituto Nacional de Investigaciones Agronómicas (INIA) by Vicente Boceta. Gallástegui’s student, Miguel Odriozola became director of
the Center of the Cuenca Alta del Ebro during the 1940s and, from 1966 until his retirement in 1973, he was professor of “zootechnology” at the Escuela Técnica Superior de Ingenieros Agrónomos (Higher Technical College of Agronomy, ETSIA) in Madrid. The CSIC was organized in three sections or “foundations,” of which the “Alonso de Herrera” was dedicated to plant biology, soil sciences, and agronomy, this was especially important because it was closely connected with the development of Spanish agriculture and with José María Albareda,105 the CSIC secretary-general, who allegedly promoted his own discipline soil sciences, to the detriment of other fields developed by
the JAE.(106)

The military uprising of 1936 had first succeeded in the agrarian and less populated areas, which were labeled “Nationalists” by Franco’s government, and did not suffer great scarcity until the end of the war, while the populated “Republican” areas experienced food scarcity from the beginning. During 1936–1939 the cereal production decreased around 30% compared to the five years’ period of 1930–1934, which is not surprising during a war. More surprising was that the Nationalist Servicio Nacional del Trigo (National Service of Wheat), established in 1937, thought to have reached a situation of wheat overproduction, so that not only was wheat production restricted, but some was also exported to Germany during Civil War. The Nationalist authorities acted as if hunger will never be their problem, but by 1939, at the end of the Civil War, Spain was overcome by food, including wheat, deficits.(107) The pre-war levels of wheat production were not reached again until the decade of the 1950s. This also happened to corn. After the war, some successful hybrids were lost and, paradoxically, the hybrid seed used in Spain since 1950s would come from France and America, due to commercial interests, while those produced by the Misión got a great success in France.(108)

The Spanish situation of political isolation afterWorldWar II – a consequence of the alignment of Francisco Franco’s dictatorship with the Axis’ Powers – and the exclusion of Spain from the Marshall Plan, marked decisively the kind of science and technology that Spain would follow in the following years.(109) Spain did not start the process of industrialization until the decade of the
1960s, remaining until that time mainly as an agricultural country. The INIA reorganization of 1940 planned the abolition of the smallest experimental stations and the regrouping of the others into major regional centers to be headed by highly reputed researchers.(110) As early as 1943, the
first meeting of applied genetics was organized in Pamplona and just after the end ofWorldWar II some important geneticists of the new period began to be trained abroad. One example, Enrique Sánchez-Monge,(111) became the most prominent agronomic geneticist in Spain. At the end of the 1940s, he took a three years fellowship abroad, first to the Sveriges Utsädesförening at Svalöf
(Sweden) in 1947 where he studied wheat genetics with Arne Müntzing, and then two years in Portugal at the Estaçâo de Melhoramento de Plantas of Elvas and the Estaçâo Agronómica Nacional of Sacavém with Antonio Souza de Câmara. During the first decade after World War II, Spain turned to Portugal as a resource, while seeking to solve the political isolation imposed
by the United Nations and many countries that would end in 1953. A strong bond was built between their agronomic institutes, with a continued exchange of students, scientists, and common projects.

Moreover, Câmara was nominated honorary director of the Laboratory of Cytogenetics of the Institute
“José Celestino Mutis,” which was established to coordinate the genetic research activities of the Spanish centers. In addition to the cooperative agreement, in 1949 the specialized journal Genética Ibérica was founded to publish the genetic research of both countries. Among the centers dedicated
to the improvement of plants, the most productive was the Experiment Station of Aula Dei, created in Zaragoza in 1950, to which Sánchez-Monge was assigned to start up a department of cytogenetics and cereal improvement between 1950 and 1957. This was a very productive period in which Joe
Hin Tjio, who determined the number of human chromosomes with Albert Levan in 1956, was part of the team. Tjio, Sánchez-Monge, and Manuel Álvarez Peña got an artificial autotetraploid of rye called “gigantón” at the end of the decade of the 1950s, but the most important achievement was the
production of an hexaploid Triticale by Tjio and Sánchez-Monge in 1954, which was improved and released for commercial production under the name of “cachirulo” in 1969.

Antoni Prevosti, considered the introducer of genetics in Catalonia during the decade of the 1950s, had a different background. Prevosti was trained as an anthropologist at the University of Barcelona, but after a visit at the Institute of Oceanography of Pallanza with Adriano Buzzati-Traverso and later at
the University of Edinburgh with Conrad Waddington, he abandoned anthropology for genetics. Prevosti had been teaching genetics at the University of Barcelona since 1955, when the old university degree of natural sciences was split into biology and geology.(112)

Prevosti initiated a research group in Drosophila subobscura population genetics in Spain, which achieved early recognition by the international community. Until 1960, no genetics chair existed in any Spanish university. Zulueta’s genetics course depended on the private chair of the Conde de Cartagena Foundation. The first university genetics chair was bestowed upon Sánchez-Monge at the ETSIA (agricultural engineering) of Madrid in 1960, which was followed closely by the one established in the Veterinary College of Zaragoza obtained by Isaías Zarazaga. In 1963 three chairs were established at the Facultad de Ciencias Biológicas (Biological Sciences School)
in Madrid, Barcelona, and Granada, which were bestowed upon Sánchez-Monge, A. Prevosti, and Eugenio Ortíz, respectively. Ortíz, also a researcher at the CSIC in the Museum of Natural Science of Madrid, left the post in Granada, which was then obtained by Amadeo Sañudo. Although Sañudo
was a natural sciences graduate, his genetics interest had developed at the Estación Agronómica para la Mejora de la Patata (Agronomic Station for Potato Improvement) in Vitoria. Years later he obtained the genetics chair of the Biological Sciences School at the Universidad Autónoma of Madrid. These four scientists completed the first generation of professors of genetics
at Spanish universities.(113)

Conclusions
Spain can be regarded as a peripheral country, as far as the establishment of genetics before and after the war is concerned. But the pattern of introduction of Morgan’s genetics is, in some respect, similar to that described by Garland Allen for Norway, Sweden, and Denmark and to a lesser degree
for Great Britain, Germany, and USSR. In those countries, the introduction of genetics depended on the existence of a cytological tradition coupled with the presence of a scientist with direct experience at Morgan’s laboratory. (114) Similarly, Zulueta’s laboratory began as a cytological one and one
of his students, Nonídez, imported the new science to Spain. The JAE had, indeed, proposed this pattern of using trainees abroad as an effective way for introducing modern science in Spain.(115)

Compared to the previous period of obscurantism, the JAE opened Spain to modern 20th century science, carrying on successfully its objective to improve the scientific level of Spain by means of fellowships abroad and the creation of research centers. Thus, modern genetics entered Spain without a great delay and was widely disseminated and understood by the principal scientific figures. However, only a few of them practiced it and they were mainly agronomists and veterinarians. The possible reason for this is that, in practice, the new genetic techniques were not completely alien for animal and plant selection practitioners, spreading faster among this group than among
the biologists, who became interested in experimental genetics only after the international success of Zulueta.

In addition, the Spanish Civil War cut too early the possible development of genetics. Only 15 years passed between the introduction of genetics in Spain (1921) and the outbreak of the Civil War (1936), not enough time for genetics to take off, particularly under the conditions of financial scarcity and political instability of Spain during the first third of the 20th century. In addition, the JAE had to create a scientific infrastructure previously non-existent. Space to work and political support were necessary to create laboratories, but these factors were not sufficient by themselves: financial support, continued training and trainees, and international collaboration as well as coordination
among Spanish scientific centers was also needed. The financial aid that the Rockefeller Foundation offered to Spanish genetics could have became the key for its future development, but this was prevented by the Civil War.

Moreover, it is possible that the Rockefeller Foundation interest to promote science would have facilitated the emergence of an international network of genetics research that neither Zulueta nor Gallástegui achieved in full, although some international collaboration took place. It deserves noting,
however, that Zulueta did not realize the importance of Drosophila research for entering into the international network, in this respect, his dedication to Phytodecta restrained to a certain extent the development of genetics in Spain. No government in Spain realized the importance and the benefits
that the development of double hybrid technology could have implied for the economy. Thus, the isolation of Spanish science continued. Moreover, it could be said that science transfer is more effective when the receiving country has research groups already at work. In Spain, in many cases, the reception of new disciplines and the creation of the research centers happened
simultaneously, which implied an added delay in the introduction and spread of the discipline.

Although many countries had suffered the effect of devastating wars, in Spain its Civil War interrupted the incipient institutionalization of scientific research and impeded the broadening of international contacts, so that the gap between Spain and the developed countries increased. The CSIC substituted the JAE, trying to erase its trace, but repeated most of its policies. The
CSIC followed the same kind of science importation by means of grants for advanced training abroad and the establishment of new centers or the reorganization of those it had inherited. The new circumstances were the power that the Church had gained again and the gradual decentralization of
research.(116) Due to the political and economic isolation of Spain during the 1940s and 1950s, the scientific community did not start to flourish until the 1960s. Just before that time, genetics began to recover, mainly in the agronomic field, where before the war genetics had been more readily accepted and which, in addition, could yield immediate benefits for the starving Spain
of the postwar. Population genetics research on Drosophila also contributed to this early recovery.

Footnotes
1 Glick, 1972, 1982.
2 Post-Kantian Karl Christian Friedrich Krause’s philosophical system was introduced into
Spain in mid-19th century by Juli´an Sanz del R´ıo, then a philosophy teacher of the University of Madrid. Krausism acquired its own character in Spain, more as an ethical guide, an attitude
and a model for a way of life, guided by reason, which in time might transform individuals
and society and thus ultimately lead to a regeneration of Spain’s long-stagnant cultural level.
3 La´ın Entralgo, 1993, vol. 1, pp. 12–13.
4 The Regeneracionismo encouraged renewal the political and intellectual life of Spain to
overcome stagnation after the defeat and loss of the colonies in 1898.
5 S´anchez Ron, 1988.
6 Most notoriously, Zulueta’s laboratory, located in a small building by the Students
Resident’s gardens, removed from the other laboratories. Originally the facility was a tool
shed. There is a famous description of this laboratory in: Gal´an, 1987a, pp. 64–65.
7 Glick, 1988.
8 Chocomeli, 1911; Lecha Marzo, 1911; Borobio D´ıaz, 1911.
9 Allen, 1978, pp. 282–283.
10 Pujiula, 1921.
11 Pujiula, 1929.
12 Besteiro, 1988.
13 Zulueta’s Curriculum Vitæ, “Relaci´on de los M´eritos Cient´ıficos y Circunstancias del
Profesor Antonio de Zulueta.” Madrid, April 3, 1935, document provided by Zulueta’s son,
Carlos de Zulueta. Also see: Carbonell, 1977; Garc´ıa Mart´ınez, 1984; Valderas, 1988; Baratas
and Fern´andez, 1989a; Baratas, 1997; Zulueta, C., 1998; Pinar, 1999a.
14 Zulueta, 1908, 1910, 1911.
15 JAE, 1914, pp. 283–284.
16 Bord´as, 1914; Non´ıdez, 1914, 1915.
17 Zulueta, 1915, 1916, 1917.
18 He bred Dutch and Russian rabbit races getting the reversion to the original brown color
of wild rabbits. He also bred albino long hair and brown short hair rabbits, whose results were
used to illustrate Mendel’s “laws” in one of the papers: Zulueta, 1926.
19 At the Archives of the National Museum of Natural Sciences, hereafter cited as MNCN,
there is an unpublished Zulueta’s translation of the 5th edition of Reginald Crundall Punnett’s
Mendelism (1919). MNCN, Scientific papers, 1913–1932. “A. Zulueta, box P183.”
20 Pinar, 1999b. The glomus aorticum is a chemoreceptor for the decrease of oxygen tension
in the blood, being less sensitive to the decrease of blood pH or the increase of carbon dioxide.
21 Archive of the JAE, hereafter cited as JAE. JAE, Letter from Non´ıdez to the president of
the JAE, Madrid, February 25, 1916. Microfiche “53–198, J. F. Non´ıdez.”
22 The JAE was mainly pro-European, yet withWorldWar I the number of fellowships to go
to the United States and neutral Switzerland increased. Economic reasons and geographical
distance limited the number who actually went to the United States. Forment´ın Ib´añez and
Villegas Sanz, 1991.
23 JAE, Letter from Non´ıdez to the president of the JAE, Madrid, February 6, 1917.
Microfiche “53–198, J. F. Non´ıdez.”
24 During the mitotic reduction some autosomes and the X-chromosomes become attached
forming a structure that functions as a whole. This structure is called X-complex.
25 Non´ıdez, 1920a.
26 Wilson, 1925, pp. 140, 778–779, 899.
27 JAE, Letter from Non´ıdez to Gonzalo Jim´enez de la Espada, New York, November
24, 1917 and fellowship renewal, Madrid, September 12, 1918. Microfiche “53–198, J. F.
Non´ıdez.”
28 JAE, 1920, pp. 47–48; Non´ıdez, 1920b, and Metz and Non´ıdez, 1921.
29 Kohler, 1994, pp. 106–109; Sturtevant quotation on p. 109.
30 Boerger, 1928, p. 376.
31 Madrid: Calpe, 1923; Madrid: Espasa-Calpe, 1936, 1938, and 1946.
32 JAE, 1922, pp. 171–172.
33 Zulueta, 1925.
34 Morgan, 1926, p. 205.
35 Morgan, 1926, p. 206.
36 Gal´an, 1931.
37 The beetle Phytodecta variabilis has four distinct color types called by Zulueta: striped
(S), yellow (Y), red (R), and black (B). Crossing experiments showed that any one of them
behaved towards any other one as an allelomorph and that the type striped was recessive to
any of the others, that yellow was recessive to the other two, and red was recessive to black.
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