G.M. Darrow, The Strawberry: History, Breeding and Physiology

6
Early Breeding in Europe

PREVIOUS CHAPTERS have indicated that the modern strawberry's development first occurred in France. It was the French who introduced F. chiloensis to Europe and by pollinating it with F. virginiana produced the first varieties of F. ananassa. Antoine Nicolas Duchesne was one of the earliest and most noted authorities on strawberry classification and history, and he must be credited as the first to identify F. chiloensis x F. virginiana as the origin of the modern strawberry. It was the English, however, who first produced the magnificent varieties of F. ananassa which started strawberry breeding in Europe and America. This English breeding work, in itself, is behind hundreds of varieties we enjoy today. After the French had seen the success of the English breeders they too began to develop their own varieties, but for half a century the "Fraises anglaises" such as Keens Seedling, the Downton, and the Elton dominated the markets of Europe and also provided a foundation for a vigorous strawberry industry in England.

The Strawberry in England
In the late 1700's and early 1800's, varieties F. virginiana Scarlet strawberry -- were the popular English garden strawberries. Attempts by English gardeners to raise new varieties from seed increased the number of varieties of F. virginiana from three to nearly thirty. James Barnet, undergardener for the Royal Horticultural Society, knew twenty-six varieties of the Scarlet in 1824. In quality most were markedly superior to the general type of Scarlets -- but none surpassed some of the wild virginiana in flavor. In size the berries showed but little improvement over the best wild ones. Growers who found that raising new plants from seed sometimes produced variations superior to the parent type, conducted experiments with seeds directly imported from Canada and North America, instead of working with American varieties imported earlier. As E. A. Bunyard observed, though slight differences resulted, the variation of F. virginiana was "not more than might have been in the wild state."1

What were these varieties? In 1766, Duchesne noted a kind of Scarlet from Strasbourg which had somewhat elongated berries; and sometime before 1826, Mr. Gibbs, a nurseryman of Old Brompton in England, raised a similar variety called the Oblong Scarlet. William Atkinson introduced the Grove End Scarlet, which he originated in 1820 at Grove End, Marylebone. Other well-known Scarlets were the Duke of Kent's Scarlet, Knight's Large Scarlet, Wilmot's Late Scarlet, Common Scarlet, and the Austrian Scarlet. The Scarlet was preferred in jam making because of its acid flavor, retention of shape, fine aroma, and bright red color. It was also the earliest strawberry to fruit naturally, without forcing.

A few varieties of pure F. chiloensis also existed in England at this time. In 1824 Barnet knew of two, possibly three, kinds, such as the Yellow Chile, which differed, mainly in the color of its fruit, from the original Chilean introduced a century earlier. F. chiloensis was unpopular in England because under most conditions it did not blossom and the fruit it did bear was poor-colored, and poor-textured, and often had a mawkish flavor. William Cobbett wrote of it: "As to the Chili it is very little superior in flavor to the potato."2

Furthermore, the Chilean was not at all hardy and was difficult to grow inland (the reason for its failure in France except at Brest on the seacoast). Philip Miller commented on its delicate constitution back in the 1750's, complaining that protection against winter frost was a precaution "absolutely necessary to the Chile strawberry, which is frequently killed in hard winters, where they are exposed without any covering." He recommended covering the beds with tanners bark, sawdust, sea coal ashes, decayed leaves, or evergreen boughs to preserve them.3 This lack of hardiness, bred into cultivated varieties, has been a major failing throughout the history of the modern strawberry.

The success of English horticulturists in producing variations of F. virginiana encouraged them in their efforts in breeding. They discovered that outstanding progress could be made when the neglected F. chiloensis was used as one parent, for these new hybrids of F. virginiana x F. chiloensis inherited characters of hardiness, sharp flavor, and high color from the Scarlet and of large fruit size from the Chile.

Behind the early, famous English hybrids lies the work of two men. The value of their contribution is seldom fully recognized, yet it is of greatest importance, for in one case a variety was produced which enters into practically all those of today, and, in the other case, breeding techniques were developed which are basic to much of the progress we now accept so casually. The first of these two men was Thomas Andrew Knight (Fig. 6-1). Although he became renowned as president of the Royal Horticultural Society from 1811 to 1838, his notable contributions to strawberry breeding have been little appreciated. Knight has been described as "the foremost pioneer in this breeding work,"4 and as "a practical breeder of new hybrids in all the important vegetables and fruits must have done more to raise the standards of excellence of these crops than any other contributor in this country before or since."5

Knight's father was a Herefordshire clergyman who died when his son was five years old. The boy's education was neglected, and until he was nine he remained almost illiterate. Since he was unable to read as a child, he concentrated his curiosity on the plant and animal life on the family estate. One day, says a story, he saw a gardener planting beans. The boy asked why the man was planting sticks of wood and was told they would grow up to be beans. The gardener's prediction came true. Knight immediately planted his pocket knife and waited in anticipation for the miraculous growth of new knives. When the experiment failed he sat down to consider the difference in the two cases. Already he was engrossed with the mysteries of the vital processes in plants, a preoccupation which would lead later to his reputation as a brilliant plant physiologist.

Eventually he did catch up with his education and with his phenomenal memory he was able to graduate from Baliol College, Oxford, with a minimum of application. He then withdrew to country life, to his little farm and greenhouse at Elton, where he would have remained a modest experimenter and breeder had not Sir Joseph Banks summoned him out of hiding. Knight first met Banks, who was then president of the Royal Society, in 1795. In April of that year Knight read his first paper before the Society on "The Grafting of Fruit Trees." Banks was interested in his ideas and began writing to him in July of 1796. Their correspondence lasted twenty years. Knight was painfully shy and reserved, but before the interest of such a great man he forgot himself and paid close attention to Banks's suggestions and advice. For some years Knight refused to read the papers of other scientists on vegetable physiology, afraid of becoming prejudiced by them in his search for truth. Consequently, he had little idea that any of his experiments and conclusions were unknown to other men. Banks finally persuaded him to read reports of other research and to publish his own ideas. He invited Knight to his house, and thus gave him the opportunity to compare his observations and theories with those of some of the world's famous naturalists. In these early years Knight was already a general farmer, stock breeder, and fruit and vegetable grower. Practicality, which was always the aim of his research, gave him an interest in improving plants by crossing them, a very new idea in 1798. From this interest came his calling as a practical agriculturist and he was included among the founders of the Royal Horticultural Society in 1804. When a committee asked him to help define the objects of the Society, he said the chief need was for "scientific inquiry into the breeding, cultivation, and forcing of fruit ... in which the practice of the modern gardener is conceived to be most defective!"6

Along this line he made the first scientific attempt at strawberry breeding on a large scale. Observing that all available varieties and species of large-fruited strawberries (with the exception of the musky one, F. moschata) would breed together, he considered them all to be varieties of the same species. From experimental crosses he raised over four hundred seedlings in 1817 in an attempt to disprove the common opinion that all European gardens were indebted to America for "three distinct species of esculent strawberry: the Grandiflora or Pine; the Chiloensis or Chile; and the Virginiana or common Scarlet."7 From the 400 seedlings he made a number of selections. He sent runners of the most promising kinds to the Royal Horticultural Society's garden in London. Each selection was numbered and had an accompanying description and account of its parentage. "I possess at present only a single bearing plant of each of the above mentioned varieties," he wrote, "but should the fruit of any be found valuable, I shall be prepared to send a very large number of plants of all . . . and such will be much in the service of every Member of the H. S., who may wish to obtain them."8

One outstanding seedling became the Downton strawberry (Plate 6-1), whose mother was raised from seed directly imported from America. Its geographical source in America was unknown, but it was classed as a Virginian. The father was the Old Black, a strawberry of unknown origin. This uncertainty about the source of the Downton's parents prevented Knight from claiming it to be a definite example of F. virginiana x F. chiloensis. The Downton itself had large oblong fruits with many resemblances to the Chilean.

The second of Knight's varieties to win attention was the Elton (Plate 6-2), which in 1828 became the Downton's rival. Knight had overlooked it in his plantings and it remained unnoticed until someone "discovered" it in the Royal Horticultural Society's garden where it was prominent for its late blooming, its beautiful fruit, its health and its hardiness. It established itself as a good late variety, but its actual parentage was unknown.

Knight's success in improving the quality of fruits and vegetables encouraged other gardeners to experiment on their own and they found Knight ready both to applaud their successes and to abet their attempts. At the end of each meeting of the Horticultural Society the members voted on the most deserving paper read at the session, which was then printed, to be preserved along with the record of the Society's transactions. By 1810, forty-one papers had been printed, fifteen of which belonged to the most prolific contributor -- Thomas Andrew Knight. The first president of the Society died in 1810 and in 1811 Knight was elected to fill his place, a position to which he was annually re-elected until his death 27 years later.

In 1806 Knight's older brother left him Downton Castle, the 10,000-acre family estate in Herefordshire. Here Knight spent the remainder of his life, journeying to London every spring for the meetings of the Society, but devoting most of his time to running the estate and its orchards and giving many hours of thought to planning his experiments.

His contributions to practical agriculture show a wide range of interests and help to explain his influence in leading other gardeners and livestock breeders to imitate his efforts. Some of the observations he made are now a standard part of plant physiology, for he made studies on the ascent and descent of sap in plants, the nature of the cambium, geotropism of roots and stems, and phototropism in tendrils. In practical horticulture "the originality and range of his investigations are practically staggering."9

In recognition of Knight's reputation and his broad knowledge, Sir Humphrey Davy, when asked to deliver a course of lectures on "The Chemistry of Agriculture" before the English Board of Agriculture in 1803, he went first, on Sir Joseph Banks's recommendation, to consult Knight on some of the points in his presentation. In 1806 Knight won the Copley medal for his papers on vegetable physiology. A close friendship developed between Knight and Davy, and in writing to him about plant physiology in 1810, Davy said: "In considering the physiology of the subject, I shall have little to do but to record your labours, for you have created almost all the science we possess upon that interesting subject."10

In 1814, Knight was awarded a gold medal by the Royal Horticultural Society for his papers as well as his gifts of grafts and buds of valuable new fruits. In 1835 a Knightian medal was struck in gold and awarded him "for signal services he has rendered to horticulture by his physiological researches."11 Knight accepted on condition he could give a sum equal to the cost of the medal to help liquidate the Society's debts.

As the years passed, Knight spent more of his time at Downton and shortened his visits to London. The death of his only son in a shooting accident in 1827 only made him work harder. He corresponded with horticultural societies and plant breeders all over Europe and America and received medals from many parts of the world. Despite his retirement to Downton, he never stopped contributing to the Society. He died in his coach on the way to the General Meeting of the Horticultural Society, May 11, 1838. As one historian said, "It would be difficult to find any other contemporary author, in this or other countries, who has made such important additions to the knowledge of horticulture and the economy of vegetation."12

Knight wanted the chief emphasis of the Royal Horticultural Society put on the breeding, cultivation and the forcing of fruits. He used his influence, first as a founding member, then as president of the Society, to insist on the awarding of medals, one of the earliest means adopted by the Society to promote horticulture. Papers and pertinent correspondence were published along with paintings of exceptional new varieties of fruits and vegetables, and silver cups went to their breeders.

The stimulus of the Royal Horticultural Society had its effect on strawberries. Michael Keens, a market gardener of Isleworth near London, raised the first large-fruited market variety of strawberry in 1806, eleven years before Knight began his crossing work, and he found the Society both interested and appreciative of his efforts. Keens became second only to Knight in his importance to the English development of the strawberry.

Having frequently, in the course of many years' practice, observed the deterioration of several kinds of fruit, when propagated in the usual ways of slips, buds, cuttings, scions, or division of the parent root, I have for a considerable time employed myself in raising new varieties from seed, which has been not only a source of great amusement to me, but also very profitable in my profession.

Keens wrote to the Society from Isleworth on January 10, 1814. He then described how he had sown the seed of the White Chili strawberry, along with a great many others in 1806 and had discovered that one of the Chili seedlings was very different from, and far superior to, its companions. This was the Keens Imperial (Plate 6-3).

In his letter to the Society, Keens described his berry as a very fine deep crimson in color and round in shape. The seeds projected above the surface of the fruit thus protecting it from bruises and suiting it for transport to the markets. Its growth was free and vigorous and its stalk was exceptionally erect and strong, better able to support the fruit than any other strawberry, which alone would give it a decided superiority over others in wet weather," Keens pointed out.13 Although it lacked a high flavor, the great beauty of its fruit and its probable usefulness for the London market persuaded the Society to publish a drawing of it.

In 1821 the Society, had its artist paint another Keens variety, the Keens Seedling (Plate 6-4), raised in 1819 from the seed of Keens Imperial. The Society awarded Keens a silver cup and published a description of the berry in "Notices of New or Remarkable Varieties of Fruits, ripened in the summer and autumn of the year 1821," a review of fruits which had been exhibited at recent meetings. Thus Knight's proposed honors went to the breeder of a strawberry which is an ancestor of most of today's leading varieties.

The berries of Keens Seedling were large for the 1820's -- up to two inches in diameter, one and a half inches in length and somewhat coxcomb in shape. The fruit was a very deep rich red in color and white at the center, though the red color stained deep into the flesh. The flesh was tender, compact and very juicy at the center with a flavor like that of the Pine strawberry. The seeds were small, yellow and deeply imbedded and the calyx was large and partially reflexed. The plants themselves were prolific and free growers and possessed"the desirable habit of bearing their fruit high from the ground," read the description in the Transactions of the Royal Horticultural Society.14 E. A. Bunyard, a strawberry expert of the early twentieth century, said that "the large size and excellent flavor of this fruit created a sensation which probably no succeeding strawberry equalled." In 1914 he noted that while other varieties might have surpassed it in individual qualities, as a near-perfect berry it had yet to be replaced.15 Keens had raised the first modern strawberry but the Keens Seedling was more of a happy chance than anything like the result of the systematic breeding program used by Knight. Other growers, spurred on by Keens' success, would use Knight's methods in their breeding.

Keens grew his strawberries in gardens extending over more than sixty acres at Worton Lane in Isleworth, the chief of the market garden towns in the region just north of the Thames, known as the "great fruit garden" of the London markets. Michael Keens and John Wilmot, originator of Wilmot's Superb Strawberry, were the chief two gardeners in early nineteenth century Isleworth. Their strawberry plantings made the township famous as a strawberry-growing center. Most of Keens' gardens were devoted to the Keens Seedling strawberry and the Keens gooseberry. Sir Joseph Banks, the friend of Thomas Andrew Knight, also lived in Isleworth at his estate "Spring Grove," named after an apple raised by Knight (Spring Grove Codlin) and which Banks cultivated extensively. Banks also grew many strawberries and reintroduced an old abandoned practice of spreading straw under the fruit to keep it clean and to diminish the amount of evaporation from the soil, thereby reducing the necessary amount of watering. Both Keens and Wilmot received the Banksian medal from the Horticultural Society for the excellence of their fruit. Keens himself owned most of his gardens "which very considerable property he attained chiefly by industry and a judicious marriage, frankly avowing that he began the world without a shilling of capital."16 His son-in-law inherited the land when Keens died in 1835 at the age of seventy-three.

Later Varieties of the 1800's*

*This section was prepared by George M. Darrow

Many excellent varieties appeared after the success of Keens Seedling. John Williams was one of the first to succeed in breeding new varieties of strawberries, which he named after his house, Pitmaston. His Pitmaston Black Scarlet was grown even as late as 1914.

In 1840, Myatt of Deptford introduced British Queen, his most famous strawberry, and one which Bunyard considered still among the best flavored in 1914. The Eleanor (1847), Admiral Dundas (1854) and Filbert Pine were other varieties raised by him. Admiral Dundas was an enormous pale orange-colored berry with pink flesh and good flavor. Eleanor was late blooming, bright red and acid and used for forcing for its very large fruits. Myatt's seedlings are supposed to have been raised from Knight's varieties. British Queen dominated the whole strawberry market for half a century. Possibly if virus-free plants were available it would be widely grown yet for its unsurpassed flavor, even though it is not hardy. Introduced into France in 1848, it was widely grown there.

Between 1840 and 1850 the Bicton Pine was introduced by J. Barnes, a gardener at Bicton in Devonshire. Although its yellowish-white, soft fruit made it an amateur's variety, its flavor was excellent, suggestive of the Pine.

In 1854, Jucunda was introduced, a variety originated by John Salter of Hammersmith, England. It has been an important variety throughout Europe and the United States; its importance greater on the continent than in England. It was imported into the United States as early as 1858, and grown under high culture and protected with winter mulching. It was very productive and was raised there till the 1920's. It was light crimson with firm, red flesh of high flavor, late and capped the easiest of any variety, being picked without caps. It is still grown slightly for processing in Holland, but has been replaced largely by Senga Sengana. It has been crossed with others in Holland to get easy capping varieties.

In the 1860's two other famous English varieties were introduced: Sir Joseph Paxton (1862) and Dr. Hogg (1866), originated by Samuel Bradley, a gardener at Elton Manor near Nottingham. The first, Sir Joseph Paxton, was a midseason variety of brilliant glossy red and firm fruit, which made it a good market berry. It was a leading variety of England for seventy-five years, and in France was considered one of their best commercial varieties as late as 1944. Dr. Hogg was a long, somewhat flattened orange-red berry with pure white flesh, but although one of the largest of the late varieties, it was not as successful as the Paxton.

About this period, Thomas Laxton (Fig. 6-2) began the most extensive breeding of strawberries ever attempted in England. He began experiments in breeding in 1865 and conducted experiments for Charles Darwin, mainly with peas. He began introducing strawberry varieties from his work as early as 1872. His first great success and the only variety he introduced that was not a handmade cross was the Noble (1884), a seedling of Excelsior that had been planted next to the American Sharpless. Until the last few years it was a major early variety of many countries. It was notable for its earliness, its resistance to cold and to disease. Even in 1960 many acres were grown in Italy and in Scandinavia. King of the Earlies (Vicomtesse Hericart de Thury x Black Prince) was introduced in 1888. In 1892, came Laxton's other great variety, Royal Sovereign, nearly equal to Keens Seedling in significance. It was a cross of Noble x King of the Earlies. It had an American variety, Sharpless, in its ancestry. The earliness, excellent flavor, beauty, productiveness, hardiness and relatively good handling quality made it of great importance in Great Britain and on the Continent. Its weaknesses are its great susceptibility to mildew and to virus diseases. After more than seventy years, it is still raised in many parts of Europe. Thomas Laxton originated seventeen varieties himself and after his sons took over the work in the 1890's, they and the grandsons introduced forty-seven more up to 1927. Scarlet Queen, Leader, Fillbasket, the Laxton, Latest, Latest of All, Bedford Champion and Duke, were some of the more notable of the other varieties. Better than other European breeders in the nineteenth century, Laxton understood the weakness of European varieties -- they were not hardy enough and needed to be hybridized with the much hardier American varieties. In the 1890's, he told of raising at least 10,000 seedlings over the thirty-two years he had been breeding strawberries.

Breeding in France-1770 to 1900

Although the modern strawberry had its beginning in the great work of Duchesne in the 1760's and 1770's, research and breeding did not continue at the same high level in France. Except for the Chilean, grown quite extensively near Brest on the coast, the Pineapple (ananassa) was the chief variety and, for most of the time, almost the only variety grown in France from the time of Duchesne, about 1770, until after the introduction from England of Keens Seedling and other varieties in 1824 and the later introduction of the Elton. There was no Thomas Knight to stimulate hybridizing as in England, nor a Hovey as in America. Most of the breeding was just the raising of seedlings by amateurs and gardeners, with little of the systematic breeding of Thomas Laxton. We know now that most seedlings of perfect-flowered varieties are selfed seedlings and far less vigorous than the parents, so that just growing seedlings is discouraging work. Only accidental crossing by bees, or growing seedlings of pistillate varieties that had to be crossed to bear fruit, would give mostly vigorous strawberry plants. But even so, just by raising seedlings, the French amateurs produced some superior varieties that were even grown in England.

In 1846, Gabriel Pelvilain, chief gardener of the royal palace at Meudon, raised the Comte de Paris and the Princesse Royale, with its hard core, both from seed of Keens Seedling and after their introduction the culture of the Pineapple strawberry soon ceased. In the same year, J. L. Jamin, raised the Vicomtesse Hericart de Thury, also from seed of the Keens Seedling, and it was introduced in 1849. This variety had excellent flavor, glossy red, firm berries, that made it the standard for flavor and preserving for one hundred years. It is still widely known under the name Ricart, from Héricart, and might still be a great variety if free from virus.

After 1849 the varieties grown were mostly of French origin until almost the end of the nineteenth century, for in the next twenty years many fine French varieties were introduced. F. Gloede introduced the Duc de Malakoff (Chilean x British Queen) in 1854, a variety of very large size. M. Lebreton selected Marguerite in 1859, from seedlings of Sir Harry x an unnamed seedling, because of its great size. It was said to average over 1/2 oz. (15 to 20 gms.) with exceptional berries of more than 11/2 oz. (40 to 45 gms.). It had pink to white flesh and was quite acid. Dr. Nicaise of Chalons sur Marne, raised the Dr. Nicaise in 1863, another berry of largest size, up to 11/2 to 2 oz. Berger obtained the Docteur Morere (Palmyre x Duc de Malakoff), in 1867 and it was introduced in 1871 as an early forcing variety of high flavor with dark raised seeds. Later it proved notable as one parent of Mme. Mouton.

Even though there was no systematic breeding in France in much of the nineteenth century, it was the French who produced during this period two great works on the strawberry. In 1864, Leonce de Lambertye, a man who used his retirement years for the intensive study Duchesne had conducted in his youth, published Le Fraisier, sa Botanique, son Histoire, sa Culture. The other work of importance was that of Decaisne, "Le Fraisier" in Le Jardin Fruitier de Museum, which included paintings of species and varieties with accompanying descriptions by Mme. Vilmorin (Fig. 6-3, see Chap. 5).

Three more varieties by three separate breeders complete the more notable ones up to 1900's. Joseph Riffaud obtained the General Chanzy (J. Riffaud x Madame Lebreton) in 1880. It was large, long ovoid, deep brilliant red, a good-flavored early sort used for forcing. Later it was the seed parent of Fukuba of Japan. Edouard Lefort originated the Edouard Lefort, a high flavored forcing variety, in 1889 and Louis Gauthier, of Caen, his Louis Gauthier, in 1889. The Louis Gauthier was a very productive, large white, very solid, rose-tinted variety, which capped with difficulty. It often produced a second crop in the fall, but on the runner plants.

At the end of the nineteenth century, Noble from England and Sharpless from America were introduced and these two and Marguerite were the main crop varieties. Docteur Morere, Marguerite and General Chanzy were the early and forcing sorts.

1 Bunyard, E. A., "The History and Development of the Strawberry," Journal of the Royal Horticultural Society, Vol. 39, 1914, p. 545.
2 Fletcher, S.W., The Strawberry in North America New York, 1917, p. 116.
3 Miller, Philip, The Gardener's Dictionary, Sixth Edition. London, 1771, n.p.
4 Pearl, R. T., The History of the Cultivated Strawberry. Wye, England, 1928,p 6.
5 Bagenal, N. B., "Thomas Andrew Knight," Journal of the Royal Horticultural Society,Vol. LXIII. London, 1938, p. 324.
6 Simmonds, A., "The History of the Royal Horticultural Society, 1804-1954," Journal of the Royal Horticultural Society, Vol. 79, London, p. 463.
7 Bagenal, op. cit., p. 207.
8 Ibid., p. 210.
9 Ibid., p. 224.
10 Knight, Thomas Andrew, Selection from the Physiological and Horticultural Papers, Published in the Transactions of the Royal and Horticultural Societies by the Late Thomas Andrew Knight, Esq. London, 1841, p. 22.
11 Ibid., p. 32.
12 Simmonds, op. cit., p. 464.
13 Keens, Michael, Transactions of the Royal Horticultural Society, Vol.II. London, p. 102.
14 "XXXI Notices of New or Remarkable Varieties of Fruits," Transactions of the Royal Horticultural Society, Vol. V. London, p. 261.
15 Bunyard , op. cit., p. 547.
16 Bate, G. E., A Middlesex Medley. England.