Biotechnology in forest tree improvement: research directions and priorities

R.J. Haines

Russell J. Haines is with the Forest Research Centre of the Queensland Department of Forestry in Gympie, Australia. He was the recipient of the FAO Andre Mayer Research Fellowship in 1992.

Cultured nodes and axilliary meristems of Araucaria cunninghamii, a valuable plantation conifer
A rooted axillary shoot of Araucaria cunninghamii

A summary of current research into biotechnology applications for forest tree improvement and recommendations for the prioritization of research objectives in the sector: This article is based on a longer work soon to be published as an FAO Forestry Paper.

In no other field of scientific investigation today are such rapid advances being made as in plant biotechnology. Biotechnology comprises any technique that uses living organisms to make or modify a product, to improve plants or animals or to develop micro-organisms for specific uses. The public imagination has been caught by such highly publicized developments as the tomato that can be frozen and the cassava and other agricultural crops that have been genetically engineered for insect and virus resistance, and which are now in or near commercial release.

The potential benefits of plant biotechnology in forestry are perhaps even greater than in agriculture because of the possibility of gaining time in certain tree improvement processes. The challenges facing foresters regarding production or outturn, whether of wood or other products, are no less urgent than those facing agriculturists.

Tree improvement research falls into two categories: supportive research, e.g. the collection of data on reproductive biology and genetics necessary to support effective breeding; and strategic research, aimed at the development of better breeding methods. Many strategic research projects concerning biotechnology have been initiated, in the opinion of some (e.g. Sedgley and Griffin, 1989) at the expense of other urgently required tree improvement activities. Clearly, the careful prioritization of research objectives is important and biotechnologies should only be used where there is an intimate basic knowledge of the species being experimented. Nonetheless, if basic biological information and knowledge are available and if sound tree improvement programmes are in place, biotechnology can be a powerful tool. This analysis is directed towards the definition of important biotechnological research priorities in forest tree improvement.

Tree improvement priorities

The general objective of a genetic improvement programme should be the sustainable management of genetic variation to produce, identify and multiply for the operational planting of well-adapted genotypes of the desired quality. Typically, this incorporates:

· the establishment of initial populations, including species and provenance testing, as well as the development of breeding and gene conservation populations;· population improvement, frequently including recurrent cycles of selection and recombination;
· the derivation and multiplication of strains to be used operationally.

In principle, the above applies to both industrial (i.e. biologically well-known species planted on a large scale) and non-industrial species, although the practicalities differ to some extent. The current status of tree improvement and associated research trends have recently been reviewed (FAO, in press; Kanowski, 1993). Significant genetic gains are being achieved in breeding programmes for established industrial species, and the broadening of such efforts will be important. Major limitations to rapid improvement with most of the established industrial species are:

Tissue cultures in an incubation cabinet

· long generation intervals, related to poor juvenile-mature correlations (i.e. that the characteristics of young trees are not necessarily accurate indicators of those found in mature individuals) and the long juvenile phase with respect to flowering;· the low effectiveness of selection for many characters as a result of low heritability or difficulties in assessment;
· through the use of open-pollinated orchards, the exploitation of only a part of the genetic variation available.

Major research priorities for established industrial species should be the broader development of methods for the propagation of full-sib families (i.e. multiple examples from a single, known male-female pair) or clones, the development of methods for early and more accurate selection and the promotion of precocious flowering. To utilize a broader range of sites, and to supply products currently provided by harvesting in natural forests, a significant proportion of new plantation areas will probably be established with tropical species that are not widely used at present. Some may be very amenable to improvement, while others present problems, e.g. flowering and seed problems and susceptibility to insects and disease. The distribution and potential uses of many species are not well known and gene pools are probably under threat. The implementation of improvement programmes will be an important priority for these species. The testing of potentially useful species, the characterization of mating systems, provenance collection, the establishment of trials on different sites, the implementation of gene conservation measures and the commencement of other breeding activities pose large challenges.

Several non-industrial taxa are highly variable and feature early and prolific flowering - conducive to very rapid improvement by traditional means. The biological features of many potentially valuable non-industrial species, however, remain largely unknown. Some gene pools are under threat. Although selection work has been undertaken in a few programmes, most non-industrial species remain at the species-testing stage. Although the genetic improvement of non-industrial species is similar in principle to that of industrial trees, and some similar limitations apply, special difficulties are presented by:

· the need to establish plantings on a wide range of sites, many of which are marginal or problematic;
· the multiplicity of selection criteria (e.g. fuelwood or mulching quality!;
· the variability of selection criteria from one grower to the next;
· the low value of forest products in some systems; and
· the difficulty of transferring breeding results to operational plantings, e.g. where growers have a strong economic preference for raising their own planting stock.

A rejuvenated shoot from an old sequoia tree

The work involved even in selecting the most promising species is formidable and, for reasons presented above, improvement beyond species and provenance testing may be difficult to justify for many non-industrial trees. Major priorities for the improvement of non-industrial species are likely to be taxonomic studies of variation; species and provenance testing; the assessment of reproductive features; and conservation activities.
As pointed out by Kanowski (1993), tree improvement does not receive adequate financial or human resource inputs. This is especially true in developing countries where funding provided through national, regional, bilateral and international programmes is not sufficient to conduct the essential activities properly. The problems are further compounded by inadequate levels of training and facilities in most areas. For the non-industrial species in particular, it is these resource limitations and the variability in user requirements, rather than biological constraints' that constitute the major impediments to rapid improvement.

Biotechnologies and tree improvement

The author has reviewed the current status of biotechnology, and applications in tree improvement in detail in a work soon to be published (FAO, in press). The following paragraphs summarize the most important biotechnologies.

Cryopreservation and in vitro storage

This comprises the maintenance of cells, tissues or organs in cultures where growth is slowed (e.g. by the reduction of light, temperature or nutrients) or suspended (by immersion in liquid nitrogen). Many technical difficulties are involved, particularly in the subsequent regeneration of plants from the cultures, but recent results are generally encouraging. Regeneration from cryopreserved tissues has been induced for more than 70 plant species, including coconut, rubber, cocoa and coffee, and for several forest tree species. These results have led to hopes that the technologies may have a number of applications in tree improvement.

Gene conservation. Although increasingly used for the storage of threatened germplasm of agricultural species (Engelmann, 1991), in vitro storage and cryo-preservation have little to offer for this purpose with regard to forest trees. Gene pools of most of the established industrial species are reasonably well preserved in stands, both in situ and ex situ, and in seed stores. Undoubtedly, the gene pool of many tree species is threatened, particularly among the tropical hardwoods and non-industrial species. Distributions of these species are poorly known, as are their biological characteristics. Major impediments to the preservation of forest tree germplasm are: the inadequacy of resources for the survey and collection work that would be required before any germplasm could be stored; and the unreliability of many existing seed storage facilities. Even for the recalcitrant (hard-to-store) species, priority would be better directed to the establishment of ex situ plantings, which should facilitate urgently needed evaluations of the material. In the longer term, cryo-preservation and in vitro storage may have some application as a backup conservation strategy, but only for populations of well-surveyed, recalcitrant species.

Maintenance of juvenility. Suspension of the growth processes also implies the maintenance of the maturation state previously attained in the tissues - without any of the uncertainty associated with alternative strategies such as long-term hedging or serial propagation. Cryopreservation therefore warrants much more attention as a means of maintaining juvenility during simultaneous clonal testing and thus capturing genetic gains offered by clonal forestry with industrial species. The technology is therefore applicable mainly in cases where good breeding programmes are in place, where clonal forestry is a realistic goal and where "rejuvenation" is difficult -particularly for the conifers.

Use of molecular markers

The use of molecular markers involves the examination, using sophisticated biochemical techniques, of variations in cellular molecules such as DNA and proteins. As an alternative to traditionally measured features such as vigour, stem quality and various morphological aspects, molecular markers offer the advantages of being unaffected by the environment or the developmental stage of the plant while also being very numerous. These characteristics have led to a number of potential applications in tree improvement.

Genetic fingerprinting. The inherent characteristics of molecular markers render them much more useful than morphological traits in establishing the identity of a particular tree or tracing its genetic relationship to other trees. For example, using molecular markers, it was possible to identify each of 39 peach cultivars individually (Ballard et al., 1992). Markers have important immediate applications in supportive research for advanced breeding programmes with industrial species mainly for quality control, e.g. checking clonal identification, orchard contamination and within-orchard mating patterns by "fingerprinting". Markers also have important immediate applications in supportive research for tropical hardwoods and non-industrial species, in particular for essential taxonomic studies and investigations of mating systems.

Quantification of genetic variation. Molecular markers are potentially more useful for quantifying genetic variation than are traits such as vigour and stem form, for which environmentally induced variation is frequently a confounding factor (i.e. it is not clear if traits are produced genetically or by external factors). Markers have been used to compare within- and inter-population variation in several tree species (Muller-Starck, Baradat and Bergmann, 1992). The quantification of genetic variation to aid in sampling strategies for the development of gene conservation and breeding populations of new industrial and non-industrial species is a potentially useful application of molecular markers. Markers may, however, provide underestimates of genetic variation with respect to traits (e.g. vigour and stem quality) that are more subject to evolutionary pressures and, therefore, will need to be used with caution.

Marker-assisted selection. This refers to indirect selection on the basis of markers shown to be associated with commercially important genes. Unaffected by the environment or developmental stage, markers offer the possibility of highly effective and early selection, long the hope of forest tree breeders (e.g. selection for wood quality at the young seedling stage). Although the possibilities are very attractive, there are limitations that will prohibit application in the short or medium term (Strauss, Lande and Namkoong, 1992): i) marker analysis is currently too expensive to permit the screening of large populations of seedlings; ii) associations between markers and economically important traits have to be established separately for different families, thus, even when cheaper markers are available, marker-assisted selection will apply mainly to advanced and sophisticated breeding programmes - those for which the creation and maintenance of the appropriate pedigree structures can be afforded and where clonal forestry is achievable. For most species, current resources would be far better directed towards moving breeding programmes to this stage of advancement, rather than to the development of marker-assisted selection.

The major current value of molecular markers lies in long-term strategic research; marker studies are making great contributions to advances in the understanding of basic genetic mechanisms and genome organization at the molecular level. An important emphasis of this work in coming years will be the study of quantitative traits of forest trees, of which a few model species will receive most attention, for example loblolly pine (Pinus taeda).

Genetic engineering

This comprises the insertion of novel genes into a plant or else the modification of existing genes through manipulation of the DNA molecule. Crops to which genes for insect, virus and selected herbicide resistance has been added are being or are near to being applied commercially. A tree crop into which these genes have been inserted is the poplar. Many projects are under way for forest trees, the reduction of lignin biosynthesis, for instance, but numerous technical difficulties remain to be solved. The insertion of currently available insect- or herbicide-resistant genes into a new species would constitute a major research undertaking, and successful application would be dependent on being able to regenerate from the transformed cells. The manipulation of more complex traits would be an even more formidable undertaking and much research remains to be done. An often overlooked research component is the extensive testing that would be required before a responsible recommendation for the large-scale deployment of transgenic plants could be made. Research projects of this type are necessarily intensive and must be regarded as long-term with only a modest expectation of success.

Insect resistance is of potential value, for example in poplars and some tropical hardwoods. However, the work involved in introducing several different resistance genes, sufficient to ensure that insects do not acquire tolerance during the rotation, should not be underestimated. The reduction of lignin biosynthesis is a very valuable objective for the pulp species. The introduction of herbicide-tolerant genes is of some interest but, in many programmes, the advantage of practicing unguarded herbicide applications may not be sufficient to pay for the research programme. Cold-tolerant genes are likely to be of some commercial value for many species, in particular the eucalypts. Much remains to be done, however, to establish that sufficient tolerance can be conferred using antifreeze proteins and to extend the work to tree species.

Prevention of the escape of genes into wild populations is likely to become an important concern, and sterility should be an early target of genetic engineering work with forest tree species. The major factor limiting application of genetic engineering in forest trees is the state of knowledge of molecular control of the traits that are of most interest - those relating to growth, adaptation and stem and wood quality. Genetic engineering of these traits remains a distant prospect.
It is important that genetically engineered genotypes be of high quality with respect to other traits as well. The clonal test is the most logical basis for the integration of genetic engineering into traditional tree improvement programmes. For these reasons, genetic engineering is most appropriately conducted with species for which breeding programmes are advanced and clonal forestry can be realistically contemplated. Research on this subject should not assume a high priority with species for which natural variation available within the taxon remains poorly investigated

Micropropagation

This refers to in vitro plant propagation methods. The principal approaches are axillary budding (actually a miniaturization of propagation with cuttings): the induction of adventitious buds on non-meristematic tissue (i.e. inducing a shoot where one would not normally develop): and somatic embryogenesis (where individual cultured cells or small groups of cells undergo development resembling that of the zygotic embryo). As an alternative to other vegetative propagation methods, the attraction of micropropagation lies in its ability to multiply elite clonal material very rapidly. More than 1 000 plant species have been micropropagated, including more than 100 forest tree species (Bajaj. 1991; Thorpe, Harry and Kumar, 1991). Successful experimental practices probably could be developed for most tree species.

For most industrial forest plantation species, the costs of planting stock and insufficient data regarding field performance remain major obstacles to be overcome before a broader use of micropropagules as direct planting stock may be contemplated (Haines, 1992). Micropropagation has an immediate application, however, in integrated clonal propagation systems featuring the commercial planting of cuttings harvested from rapidly multiplied, micropropagated stool plants of the selected clones. This approach is of value only in very advanced breeding programmes incorporating the identification of outstanding clones currently only a few programmes are at this level. Appropriate integration into breeding programmes is essential. Where clonal testing on a relatively large scale is possible and affordable, the current applicability of techniques mainly to juvenile material is not necessarily an impediment to the capture of good gains through clonal forestry. This conclusion, however, is dependent on the ability to store juvenile material for the period of a clonal test. Genetic variation in response, often substantial, is not likely to be a major problem where clonal testing can be preceded by screening for responsive genotypes, although demonstrating the absence of adverse correlations with economic traits is important. Breeding programmes new industrial species and non-industrials are not sufficiently advanced to warrant much use of micropropagation in the short term.

Field testing of loblolly pine (Pinus taeda) tissue culture plants and Dings of the same family. Plantlets display slightly more mature features

Micropropagation may have a wider application in the multiplication of stool plants of industrial species as breeding programmes become more advanced and other limitations to clonal forestry (e.g. maturation problems) are overcome. For some non-industrial tree species, micropropagation may ultimately have a role in the multiplication of selected varieties prior to release. Development of simple micropropagation techniques for those species for which such methods are not already available is therefore a useful research objective but should not take priority over issues such as advancement of the breeding programme.

Work done with some crop species indicates the possibility of encapsulating somatic embryos to form artificial seeds which can then be handled like conventional seeds. With considerable research, developments in this area may overcome the constraint of planting stock costs (discussed above) and enable the direct use of such propagules in forest plantation establishment. For industrial species, therefore, the development of these technologies is a useful long-term research objective but one which is best pursued with one or two model species, for example Picea abies and Pinus taeda.

In vitro control of the maturation state

There have been several reports of cultured mature buds displaying a reversion to a more juvenile state in response to the culture techniques and conditions. This has led to hopes that in vitro rejuvenation may be the solution to the poor rooting and vigour displayed by shoots collected from trees of selectable age for many forest plantation species. The major limitation to this approach is that there is little evidence of complete, permanent and reliable rejuvenation. In fact, some studies have clearly demonstrated the effect to be a temporary response to the culture condition. Further empirical work with this objective has a low probability of success. An understanding of the molecular basis of maturation (e.g. Hutchison and Greenwood, 1991) is much more likely to lead to practical manipulation, but this work is in its infancy and the reversal or promotion of maturation to precise levels remains a distant prospect.
For clonal forestry with industrial species, the maintenance of juvenility is about as useful as rejuvenation for many purposes (Haines, 1992) and it is probably able to be achieved using technologies such as cryopreservation or coppicing. Nevertheless, a more fundamental control of the maturation state remains one of the most valuable objectives of long-term strategic research in forest tree improvement with industrial species. Rejuvenation is most applicable to efforts where good breeding programmes are in place and where other limitations to clonal forestry do not exist.

Research Priorities

Supportive research

The preceding analysis suggests that the short-term possibilities for applying biotechnology in supportive research in forestry are:

· the use of molecular markers for quality control in advanced breeding programmes with established industrial species, e.g. for checking clonal identification, orchard contamination and within-orchard mating patterns by fingerprinting;· the use of markers in essential taxonomic studies and investigations of mating systems;
· the use of markers for the quantification of genetic variation to aid in the design of sampling strategies for gene conservation and breeding population collections for breeding programmes with "new" industrial and non-industrial species.

Strategic Research

Strategic research priorities relating to the application of biotechnology in tree improvement can be grouped into three broad areas:
Long-term generic research. This is most efficient if conducted collaboratively with a small number of model species, thus avoiding the diffusion of resources and efforts. High priority should be accorded to:

· genetic engineering for sterility - this will underlie many of the eventual applications of genetic engineering;· the use of molecular markers and DNA transformation techniques to investigate genetic processes at the molecular level, in particular those relating to complex traits such as growth, adaptation and stem and wood quality, is of particular relevance to industrial species, but will also pave the way for the application of biotechnology to non-industrial trees;
· molecular studies of the maturation state for industrial plantation species.

A somewhat lower priority should be given to the development of somatic embryogenesis in combination with artificial seed technology as an inexpensive method of clonal propagation.
Long-term specific research. Two high priority areas are:

· genetic engineering of useful traits, including lignin reduction in pulp species; cold tolerance, particularly in eucalypts; and insect resistance, e.g. in poplars and perhaps Meliaceae (when appropriate breeding programmes are in place). Transformation with appropriate genes (the introduction of several genes in the case of insect resistance) may be achieved within the short to medium term (the next five to ten years) but must be followed by perhaps ten years of field testing before responsible commercial deployment may be recommended;· marker-assisted selection, for species where breeding is advanced and where the creation and maintenance of the appropriate population structures are feasible and affordable - it will probably be ten years before this is possible on an operational scale.

Short- to medium-term research. Areas that warrant attention include:

· the examination of genetic correlations between regenerative competence and commercially important field traits (high priority);· the development of cryopreservation methods as a means of maintaining juvenility in advanced breeding programmes with industrial species (high priority);
· the development of cryopreservation as a backup measure for gene conservation in proven species for which breeding programmes are in existence and for which seed recalcitrance has been demonstrated (moderate priority);
· the development of simple micropropagation techniques for species where none is yet available (low to moderate priority).

Conclusion

Modern biotechnology should be perceived as a new group of tools or means to be used as adjuncts or complements to conventional technologies in solving problems and meeting the needs of human beings. A balance should be maintained between modern biotechnological and conventional research, and the development and application of biotechnology should be driven by needs and not by technological capability. The use of modern biotechnologies should be promoted for more efficient solutions to problems already on the agenda and within the framework of the existing priorities of individual countries. Thus, the funding of biotechnological research initiatives cannot and must not be at the expense of conventional genetic improvement programmes.

Bibliography

Bajaj, Y.P.S. 1991. Automated micropropagation for en masse production of plants. In Y.P.S. Bajaj, ed. Biotechnology in agriculture and forestry 17 High-tech and micropropagation 1, p. 3- 16. Berlin, Springer Verlag.
Ballard, R.E., He, G., Abott, A.G., Mink, G. & Belthoff, L.E. 1992 Molecular biology of forest trees. DNA fingerprinting of Prunus varieties using low copy sequence probes. Proc. IUFRO Working Party S2.04.06 Workshop. Carcans-Maubuisson France. INRA.
Engelmann, F. 1991. In vitro conservation of tropical plant germplasm - a review. Euphytica, 57: 227-243.
FAO. The role of biotechnology in forest tree improvement, with particular reference to developing countries. FAO Forestry Paper No. 118. (in press)
Haines, R.J. 1992. Mass production technology for genetically improved, fast growing forest tree species. Mass propagation by cuttings, biotechnologies, and the capture of genetic gain. Paper presented at IUFRO symposium. Bordeaux, France.
Hutchison, K.W. & Greenwood, M.S. 1991. Molecular approaches to gene expression during conifer development and maturation. Ecol. Manage.. 43: 273-286.
Kanowski, P.J. 1993. Forest genetics and tree breeding. Plant Breed. Abstr., 63: 717-726.
Mather, A. 1990. Global forest resources. London Belhaven.
Moran, G.F. 1992. Patterns of genetic diversity in Australian tree species. New Forests. 6: 49-66.
Muller-Starck, G., Baradat, P. & Bergmann, F. 1992. Genetic variation within European tree species. New Forests 6: 23-47.
Sedgley, M. & Griffin, A.R. 1989. Sexual reproduction of tree crops. London. Academic.
Strauss, S.H., Lande, R. & Namkoong, G. 1992. Limitations of molecular-marker-aided selection in forest tree breeding. Can. For. Res., 22: 1050-1061.
Thorpe, T.A., Harry, I.S. & Kumar, P.P. 1991. Application of micropropagation to forestry. In P.C. Debergh & P.H. Zimmerman, eds. Micropropagation: technology and application. Dordrecht, the Netherlands Kluwer Academic.
Weedon, N.F. 1989. Applications of isozymes in plant breeding. Plant Breed Rev., 6: 11-54.

For further information log on website:

http://www.fao.org/docrep/t2230e/t2230e0a.htm

COTTAGE GARDEN

The cottage garden is a distinct style of garden that uses an informal design, traditional materials, dense plantings, and a mixture of ornamental and edible plants. English in origin, the cottage garden depends on grace and charm rather than grandeur and formal structure. Homely and functional gardens connected to working-class cottages go back several centuries, but their reinvention in stylised versions grew in 1870s England, in reaction to the more structured and rigorously maintained English estate gardens that used formal designs and mass plantings of brilliant greenhouse annuals.

Roses, clematis, a thatched roof: a cottage garden in Brittany.

The earliest cottage gardens were more practical than their modern descendants — with an emphasis on vegetables and herbs, along with some fruit trees, perhaps a beehive, and even livestock. Flowers were used to fill any spaces in between. Over time, flowers became more dominant. The traditional cottage garden was usually enclosed, perhaps with a rose-bowered gateway. Flowers common to early cottage gardens included traditional florist's flowers, such as primroses and violets, along with flowers chosen for household use, such as calendula and various herbs. Others were the old-fashioned roses that bloomed once a year with rich scents, simple flowers like daisies, and flowering herbs. Over time, even large estate gardens had sections they called "cottage gardens".

Modern-day cottage gardens include countless regional and personal variations of the more traditional English cottage garden, and embrace plant materials, such as ornamental grasses or native plants, that were never seen in the rural gardens of cottagers. Traditional roses, with their full fragrance and lush foliage, continue to be a cottage garden mainstay — along with modern disease-resistant varieties that keep the traditional attributes. Informal climbing plants, whether traditional or modern hybrids, are also a common cottage garden plant. Self-sowing annuals and freely spreading perennials continue to find a place in the modern cottage garden, just as they did in the traditional cottager's garden.

History

Vernacular thatched cottages (built in 1812–1816) in Woburn Street, Ampthill, Bedfordshire, surrounded by garden.

Origins

Cottage gardens, which emerged in Elizabethan times, appear to have originated as a local source for herbs and fruits. One theory is that they arose out of the Black Death of the 1340s, when the death of so many laborers made land available for small cottages with personal gardens. According to the late 19th-century legend of origin, these gardens were originally created by the workers that lived in the cottages of the villages, to provide them with food and herbs, with flowers planted in for decoration. Helen Leach analysed the historical origins of the romanticised cottage garden, subjecting the garden style to rigorous historical analysis, along with the ornamental potager and the herb garden. She concluded that their origins were less in workingmen's gardens in the 19th century and more in the leisured classes' discovery of simple hardy plants, in part through the writings of John Claudius Loudon. Loudon helped to design the estate at Great Tew, Oxfordshire, where farm workers were provided with cottages that had architectural quality set in a small garden—about an acre—where they could grow food and keep pigs and chickens.

Authentic gardens of the yeoman cottager would have included a beehive and livestock, and frequently a pig and sty, along with a well. The peasant cottager of medieval times was more interested in meat than flowers, with herbs grown for medicinal use and cooking, rather than for their beauty. By Elizabethan times there was more prosperity, and thus more room to grow flowers. Even the early cottage garden flowers typically had their practical use—violets were spread on the floor (for their pleasant scent and keeping out vermin); calendulas and primroses were both attractive and used in cooking. Others, such as sweet william and hollyhocks were grown entirely for their beauty.

Dévelopment

The "naturalness" of informal design began to be noticed and developed by the British leisured class. Alexander Pope was an early proponent of less formal gardens, calling in a 1713 article for gardens with the "amiable simplicity of unadorned nature". Other writers in the 18th century who encouraged less formal, and more natural, gardens included Joseph Addison and Lord Shaftesbury. The evolution of cottage gardens can be followed in the issues of The Cottage Gardener (1848–61), edited by George William Johnson, where the emphasis is squarely on the "florist's flowers", carnations and auriculas in fancy varieties that were originally cultivated as a highly competitive blue-collar hobby.

Restored Gertrude Jekyll border at Manor House, Upton Grey, Hampshire.

William Robinson and Gertrude Jekyll helped to popularise less formal gardens in their many books and magazine articles. Robinson's The Wild Garden, published in 1870, contained in the first edition an essay on "The Garden of British Wild Flowers", which was eliminated from later editions. In his The English Flower Garden, illustrated with cottage gardens from Somerset, Kent and Surrey, he remarked, "One lesson of these little gardens, that are so pretty, is that one can get good effects from simple materials." From the 1890s his lifelong friend Jekyll applied cottage garden principles to more structured designs in even quite large country houses. Her Colour in the Flower Garden (1908) is still in print today.

Robinson and Jekyll were part of the Arts and Crafts Movement, a broader movement in art, architecture, and crafts during the late 19th century which advocated a return to the informal planting style derived as much from the Romantic tradition as from the actual English cottage garden. The Arts and Crafts Exhibition of 1888 began a movement toward an idealised natural country garden style. The garden designs of Robinson and Jekyll were often associated with Arts and Crafts style houses. Both were influenced by William Morris one of the leaders of the Arts and Crafts Movement—Robinson quoted Morris's views condemning carpet bedding; Jekyll shared Morris's mystical view of nature and drew on the floral designs in his textiles for her gardening style. When Morris built his Red House in Kent, it influenced new ideas in architecture and gardening—the "old-fashioned" garden suddenly became a fashion accessory among the British artistic middle class, and the cottage garden esthetic began to emigrate to America.

Modern Day

In the early 20th century the term "cottage garden" might be applied even to as large and sophisticated a garden as Hidcote Manor, which Vita Sackville-West described as "a cottage garden on the most glorified scale" but where the colour harmonies were carefully contrived and controlled, as in the famous "Red Borders". Sackville-West had taken similar models for her own "cottage garden", one of many "garden rooms" at Sissinghurst Castle - her idea of a cottage garden was a place where "the plants grow in a jumble, flowering shrubs mingled with Roses, herbaceous plants with bulbous subjects, climbers scrambling over hedges, seedlings coming up wherever they have chosen to sow themselves". The cottage garden ideal was also spread by artists such as water-colourist Helen Allingham (1848–1926). Another influence was Margery Fish (1892–1969), whose garden survives at East Lambrook Manor.

Helen Allingham watercolor showing elite peonies and modern delphiniums in an idealised cottage garden.

The cottage garden in France is a development of the early 20th century. Monet's garden at Giverny is a prominent example, a sprawling garden full of varied plantings, rich colors, and water gardens. In modern times, the term 'cottage garden' is used to describe any number of informal garden styles, using design and plants very different from their traditional English cottage garden origins. Examples include regional variations using a grass prairie scheme (in the American midwest) and California chaparral cottage gardens.

Design

While the classic cottage garden is built around a cottage, many cottage-style gardens are created around houses and even estates such as Hidcote Manor, with its more intimate "garden rooms". The cottage garden design is based more on principles than formulae: it has an informal look, with a seemingly casual mixture of flowers, herbs, and vegetables often packed into a small area. In spite of their appearances, cottage gardens have a design and formality that help give them their grace and charm. Due to space limitations, they are often in small rectangular plots, with practical functioning paths and hedges or fences. The plants, layout, and materials are chosen to give the impression of casualness and a country feel. Modern cottage gardens frequently use local flowers and materials, rather than those of the traditional cottage garden. What they share with the tradition is the unstudied look, the use of every square inch, and a rich variety of flowers, herbs, and vegetables.

The cottage garden is designed to appear artless, rather than contrived or pretentious. Instead of artistic curves, or grand geometry, there is an artfully designed irregularity. Borders can go right up to the house, lawns are replaced with tufts of grass or flowers, and beds can be as wide as needed. Instead of the discipline of large scale color schemes, there is the simplicity of harmonious color combinations between neighbouring plants. The overall appearance can be of "a vegetable garden that has been taken over by flowers." The method of planting closely packed plants was supposed to reduce the amount of weeding and watering required, but planted stone pathways or turf paths, and clipped hedges overgrown with wayward vines, are cottage garden features requiring well-timed maintenance.

Materials

Paths, arbors, and fences use traditional or antique looking materials. Wooden fences and gates, paths covered with locally made bricks or stone, and arbors using natural materials all give a more casual—and less formal—look and feel to a cottage garden. Pots, ornaments, and furniture also use natural looking materials with traditional finishes—everything is chosen to give the impression of an old-fashioned country garden.
Plants

Cottage garden plants are chosen for their old-fashioned and informal appeal. Many modern day gardeners use heirloom or 'old-fashioned' plants and varieties—even though these may not have been authentic or traditional cottage garden plants. In addition, there are modern varieties of flowers that fit into the cottage garden look. For example, modern roses developed by David Austin have been chosen for cottage gardens because of their old-fashioned look (multi-petaled form and rosette-shaped flowers) and fragrance—combined with modern virtues of hardiness, repeat blooming, and disease-resistance. Modern cottage gardens often use native plants and those adapted to the local climate, rather than trying to force traditional English plants to grow in an incompatible environment—though many of the old favorites thrive in cottage gardens throughout the world.
Roses

Cottage gardens are always associated with roses: shrub roses, climbing roses, and old garden roses with lush foliage, in contrast to the gangly modern hybrid tea roses. Old cottage garden roses include cultivated forms of Rosa gallica, which form dense mounded shrubs 3–4 ft high and wide, with pale pink to purple flowers—with single form to full double form blooms. They are also very fragrant, and include the ancient Apothecary's rose (R. gallica 'Officinalis'), whose magenta flowers were preserved solely for their fragrance. Another old fragrant cottage garden rose is the Damask rose, which is still grown in Europe for use in perfumes. Cultivated forms of this grow 4 to 6 ft or higher, with gently arching canes that help give an informal look to a garden. Even taller generally are the Alba roses, which are not always white, and which bloom well even in partial shade.

A climbing sport of the elite 'Souvenir de la Malmaison, introduced before 1893, typical of a modern cottage garden.

The Provence rose or Rosa centifolia is the full and fat "cabbage rose" made famous by Dutch masters in their 17th-century paintings. These very fragrant shrub roses grow 5 ft tall and wide, with a floppy habit that is aided by training on an arch or pillar. The centifolia roses have produced many descendants that are also cottage garden favorites, including varieties of moss rose (roses with attractive 'mossy' growth on their flower stalks and flower buds). Unlike most modern hybrids, the older roses bloom on the previous year's wood, so they aren't pruned back severely each year. Also as they don't bloom continuously, they can share their branches with later-flowering climbers such as Clematis vines, which use the rose branches for support. A rose in the cottage garden is not segregated with other roses, with bare earth or mulch underneath', but is casually blended with other flowers, vines, and groundcover.

With the introduction of China roses (derived from Rosa chinensis) late in the 18th century, many hybrids were introduced that had the remontant (repeat-blooming) nature of the China roses, but maintained the informal old rose shape and flower. These included the Bourbon rose and the Noisette rose, which were added to the rose repertoire of the cottage garden, and, more recently, hybrid "English" roses introduced by David Austin.

Climbing Plants

Many of the old roses had cultivars that grew very long canes, which could be tied to trellises or against walls. These older varieties are called "ramblers", rather than "climbers". Climbing plants in the traditional cottage garden included European honeysuckle (Lonicera periclymenum) and Traveller's Joy (Clematis vitalba). The modern cottage garden includes many Clematis hybrids that have the old appeal, with sparse foliage that allows them to grow through roses and trees, and along fences and arbors. There are also many Clematis species used in the modern cottage garden, including Clematis armandii, Clematis chrysocoma, and Clematis flammula. Popular honeysuckles for cottage gardens include Japanese honeysuckle and Lonicera tragophylla.

Clematis vitalba.

Hedging Plants

In the traditional cottage garden, hedges served as fences on the perimeter to keep out marauding livestock and for privacy, along with other practical uses. Hawthorn leaves made a tasty snack or tea, while the flowers were used for making wine. The fast-growing Elderberry, in addition to creating a hedge, provided berries for food and wine, with the flowers being fried in batter or made into lotions and ointments. The wood had many uses, including toys, pegs, skewers, and fishing poles. Holly was another hedge plant, useful because it quickly spread and self-seeded. Privet, was also a convenient and fast-growing hedge. Over time, more ornamental and less utilitarian plants became popular cottage garden hedges, including laurel, lilac, snowberry, japonica, and others.
Flowers and Herbs

Popular flowers in the traditional cottage garden included florist's flowers which were grown by enthusiasts—such as violets, pinks, and primroses —and those grown with a more practical purpose. For example, the calendula, grown today almost entirely for its bright orange flowers, was primarily valued for eating, for adding color to butter and cheese, for adding smoothness to soups and stews, and for all kinds of healing salves and preparations. Like many old cottage garden annuals and herbs, it freely self-sowed, making it easier to grow and share. Other popular cottage garden annuals included violets, pansies, stocks and mignonette.

Lavender.

Perennials were the largest group of traditional cottage garden flowers —those with a long cottage garden history include hollyhocks, carnations, sweet williams, marguerites, marigolds, lilies,  peonies,  tulips, crocus, daisies,  foxglove, monkshood, lavender, campanulas, Solomon's seal, evening primrose, lily-of-the-valley, primrose, cowslips, and many varieties of roses.

Today herbs are typically thought of as culinary plants, but in the traditional cottage garden they were considered to be any plant with household uses. Herbs were used for medicine, toiletries, and cleaning products. Scented herbs would be spread on the floor along with rushes to cover odors. Some herbs were used for dyeing fabrics. Traditional cottage garden herbs included sage, thyme, southernwood, wormwood, catmint, feverfew,  lungwort, soapwort, hyssop, sweet woodruff, and lavender.

Fruits

Fruit in the traditional cottage garden would have included an apple and a pear, for cider and perry,  gooseberries and raspberries. The modern cottage garden includes many varieties of ornamental fruit and nut trees, such as crabapple and hazel, along with non-traditional trees like dogwood.
References

1. ^ Ryrie, Charlie (2004). The Cottage Garden: How to Plan and Plant a Garden That Grows Itself. Collins & Brown.p. 7. ISBN 1-84340-216-5.
2. ^ ^a b Scott-James, Anne; Osbert Lancaster (2004). The Pleasure Garden: An Illustrated History of British Gardening. Frances Lincoln Publishers. p. 80. ISBN 978-0-7112-2360-8.
3. ^ Anne Scott-James, The Cottage Garden (London: Lane) 1981, de-mythologised the origins of the English cottage garden, and its treasured topiary among the vegetables and flowers, popularly supposed to represent heirlooms from the seventeenth century.
4. ^ Leach, Helen (2000). Cultivating Myths: Fiction, Fact and Fashion in Garden History. Auckland: Godwit. ISBN 1-86962-049-6.
5. ^ A Biographical Dictionary of British Architects, 1600–1840, Howard Colvin, Yale University Press, 2008 ISBN 0-300-12508-9, p 659.
6. ^ Lloyd, Christopher; Richard Bird (1999). The Cottage Garden. Jacqui Hurst. Dorling Kindersley. pp. 6–9. ISBN 978-0-7513-0702-3.
7. ^ Reynolds, Myra (1896). The Treatment of Nature in English Poetry Between Pope and Wordsworth. The University of Chicago Press.p. 253.
8. ^ Gould, Jim (Spring 1988). "The Lichfield Florists". Garden History (Garden History, Vol. 16, No. 1) 16 (1): 17–23. doi:10.2307/1586902. JSTOR 1586902.
9. ^ Massingham, Betty (1978). "William Robinson: A Portrait". Garden History (The Garden History Society) 6 (1): 61–85. doi:10.2307/1586557. JSTOR 1586557. p. 63f.
10. ^ Massingham, p. 71.
11. ^ Kendle, Tony; Stephen Forbes (1997). Urban Nature Conservation: Landscape Management in the Urban Countryside. Taylor & Francis. p. 79. ISBN 978-0-419-19300-5.
12. ^ Horwood, Catherine (2008). Potted History: The Story of Plants in the Home. Frances Lincoln Publishers. p. 151. ISBN 978-0-7112-2800-9.

Bibliography

• Clayton, Virginia Tuttle (2000). The Once and Future Gardener: Garden Writing from the Golden Age of Magazines, 1900–1940. David R. Godine Publisher. ISBN 978-1-56792-102-1.
• Garland, Sarah (2003). The Herb Garden. Frances Lincoln Publishers. ISBN 978-0-7112-2057-7.
• Gould, Jim (Spring 1988). "The Lichfield Florists". Garden History (Garden History, Vol. 16, No. 1) 16 (1): 17–23. doi:10.2307/1586902.  JSTOR 1586902.
• Horwood, Catherine (2008). Potted History: The Story of Plants in the Home. Frances Lincoln Publishers. ISBN 978-0-7112-2800-9.
• Hunt, John Dixon; Joachim Wolschke-Bulmahn (1993). The Vernacular Garden: Dumbarton Oaks Colloquium on the History of Landscape Architecture XIV. Dumbarton Oaks. p. 9. ISBN 978-0-88402-201-5.

• Kammen, Carol; Norma Prendergast (2000). Encyclopedia of Local History. Rowman Altamira. ISBN 978-0-7425-0399-1.
• Kendle, Tony; Stephen Forbes (1997). Urban Nature Conservation: Landscape Management in the Urban Countryside. Taylor & Francis. ISBN 978-0-419-19300-5.
• Lloyd, Christopher; Richard Bird (1999). The Cottage Garden. Jacqui Hurst. Dorling Kindersley. ISBN 978-0-7513-0702-3.
• Massingham, Betty (1978). "William Robinson: A Portrait". Garden History (The Garden History Society) 6 (1): 61–85. doi:10.2307/1586557.  JSTOR 1586557.
• Reynolds, Myra (1896). The Treatment of Nature in English Poetry Between Pope and Wordsworth. The University of Chicago press.
• Sackville-West, "Hidcote Manor", Journal of the Royal Horticultural Society 74 (1949:476-81), noted by Brent Elliott, "Historical Revivalism in the Twentieth Century: A Brief Introduction" Garden History 28.1, Reviewing the 20th-century Landscape (Summer 2000:17–31)
• Schulman, Andrew; Jacqueline Koch (2004). The Northwest Cottage Garden. Sasquatch Books. ISBN 978-1-57061-363-0.
• Scott-James, Anne (1981). The Cottage Garden. Allen Lane. ISBN 978-0-7139-1263-0.
• Scott-James, Anne; Osbert Lancaster (2004). The Pleasure Garden: An Illustrated History of British Gardening. Frances Lincoln Publishers. ISBN 978-0-7112-2360-8.
• White, Lee Anne (2001). Exploring Garden Style: Creative Ideas from America's Best Gardeners. Taunton Press. ISBN 978-1-56158-474-1.

External Links

• Royal Horticultural Society, British gardening charity
• The Cottage Garden Society, British Gardening society

Wikipedia