Watsonias are grown as garden ornamentals for their spectacular display of flowers in spring. The wild species introduced from South Africa in the 19th century all had flowers in shades of pink, red, orange or purple. H.H. Arderne’s discovery of a pure white-flowered, acyanic, clone of Watsonia borbonica was big news in 1888, and soon it was a feature display at Kew Gardens. Its popularity and wide dissemination as a garden plant are documented by the numerous articles in gardening magazines of that time. Early 20th century breeders – notably John Cronin in Australia and Luther Burbank in the USA – used Arderne’s White with the species then available in their hybridisation and selection programs.
Acyanic Watsonia plants have white flowers with cream anthers. They also have completely green leaves, and stems, because they lack the anthocyanin pigments. On the other hand, wild-type plants have coloured flowers with purple anthers; their leaves may have brown or red pigmentation on the margins and bases.
Some results from my watsonia breeding program from 1997 to 2010 strongly suggested that Arderne’s White and five other acyanic cultivars derived from Watsonia borbonica are due to a single recessive allele, a, with the wild-type allele A allowing the formation of anthocyanins. The same allele was present in some coloured cultivars, as shown by the appearance of roughly 25% acyanic progeny when these were crossed. It also produced recessive acyanic progeny in experimental crosses with Watsonia aletroides and W. meriana. It seems most likely that the a allele was introduced into cultivation via Arderne’s White. But since it is a recessive it must have been present in the gene pool before a rare homozygous plant happened to appear and survive long enough for Mr Arderne to notice it.
All the acyanic Watsonia accessions that I examined have something else in common. Their production of anthocyanins could have been blocked at several different points in the chemical pathway, but in fact they all lack the enzyme chalcone synthase, which is needed to produce all the flavonoid pigments, the yellow chalcones as well as the anthocyanins. This fits the explanation that they all have a mutation at the same A locus that inhibits the production of this particular enzyme. To complicate matters, some temporarily show a pink tinge when the white flowers open in cold weather. The possibility that these have a complementary mutation at a different locus was ruled out by a test cross, which produced only acyanic progeny. If two loci had been involved, only cyanic progeny would have been expected from this cross.
A classic, but all too brief, paper by Horn (1963) mentioned the interesting observation of a 3:5 segregation ratio when he crossed Arderne’s White with three different Watsonia hybrids that had coloured flowers, implying that the acyanic gene was dominant. The lack of further detail makes it difficult to compare his experiment with mine. It may be that one or two of the three hybrids were Aa heterozygotes, giving some acyanic progeny when crossed with the homozygous aa Arderne’s White.
Many Watsonia cultivars have been characterised in horticultural literature and retail catalogues simply as ‘white-flowered’. However, a named cultivar of an ornamental perennial is normally a vegetatively propagated clone that has constant horticultural properties due to its constant genotype. Descriptions need to be based on several characters if we want to define unambiguously cultivars with unique genotypes. Descriptions would ideally be supported by a type specimen and molecular data as well. A decade ago there was a lot of talk about cultonomy – the taxonomy of cultivated plants – as a separate subject, and there was still an International Code of Nomenclature for Cultivated Plants separate from the more general International Code of Nomenclature used for plants, fungi and algae. We seem to have gone backwards since then, and I fear that the Procrustean attempt to fit plants created by humans into a system of nomenclature intended for wild taxa will cause a gradual loss of knowledge of cultivars.
Ornamental flower cultivars are selected for the shape and colours of their flowers, and for growth traits that make them ‘gardenable’, i.e. convenient to use in gardens. But other morphological characters that have not been directly selected may be useful indicators to distinguish related cultivars. Among Watsonia cultivars, these characters include the relative length of the style, shape of floral bracts, orientation of stamens and shape of the seed capsule. For example, two genotypes of white-flowered Watsonia that John Cronin produced in the 1920s had become lumped together in the old nursery that preserved them: both are tall slender plants with white funnel-shaped flowers. But one has a style equal equal in length to its stamens, while the other has a much longer style that protrudes from the flower. When the two are grown side by side for a few years, other small differences in flowering time, mean height and maximum leaf width can be noticed.
Cooke, D.A. (2010) Genetics of white-flowered cultivars derived from Watsonia borbonica (Iridaceae). J.Adelaide Bot. Gard. 24: 33-38. Download the PDF here
Horn, W. (1963) Flower colour inheritance in some Iridaceae. Naturwissenschaften 50(15): 527-528.
Wisdom has no great tradition in the Western world. But if we are very industrious, it will be up to us to make one.
L. Ron Hubbard Phoenix Lectures, 19 July 1954
The South African Chasmanthe floribunda (Salisb.)N.E.Br., is a common garden plant in southern Australia. It could be called a low-care relative of Gladiolus, in the family Iridaceae. The wild-type has an orange-red perianth with some yellow on the perianth tube, a purple inflorescence axis and purple anthers. But some of the plants in cultivation have a completely yellow perianth and yellow anthers, with a green inflorescence axis.
I have examined perianths of both flower types under a light microscope at 200X. The red anthocyanin pigment was dispersed through the cell sap in the red flowers, but could not be found in the yellow flowers; the yellow pigment (presumably a carotenoid) was concentrated in chromoplasts in both red and yellow flowers.
No plants with intermediate flower colour were observed, although the species regenerates freely from self-sown seed (to the extent of becoming a weed of native vegetation). Because of this clear distinction, the yellow variant had been formally named as C. floribunda var. duckittii by Louisa Bolus (1933).
Apart from its lack of red pigment, the yellow variant is indistinguishable from the wild-type in its morphology. De Vos (1985) had suggested that the portion of the perianth tube below the insertion of the stamens (the hypanthium) was shorter in the yellow flowers but did not cite any measurements. However, flowers that I have measured in suburban Adelaide gardens and roadside populations in the Adelaide hills have hypanthium lengths of 9-11 mm, irrespective of their flower colour.
The red pigment is also absent from the plumule of the seedling and etiolated shoots emerging from the corms; they are cream in this variant but reddish-tinted in wild-type plants. This suggested the simple experiment of crossing red- and yellow-flowered plants and scoring their progeny at germination for pigment.
Cross pollination was carried out between a ‘red’ and a ‘yellow’ plant; reciprocal crosses were made to check for any apomixis or accidental self-pollination.
- Flowers of a ‘red’ were pollinated with pollen from a ‘yellow’
Result: 30 ‘red’ seedlings, 32 ‘yellow’
- Flowers of a ‘yellow’ were pollinated with pollen from a ‘red’
Result: 31 ‘red’ seedlings, 35 ‘yellow’
These results are in accord with a 1:1 ratio (Chi-square 0.281, yielding a probability of 0.595). Four randomly selected plants from each of the ‘red’ and ‘yellow’ seedlings were grown through to flowering; all four scored as red produced wild-type red flowers, all those scored as yellow produced pure yellow. No intermediates between red and yellow were produced in the experiment.
The simplest hypothesis is that a single gene is involved, with red fully dominant over yellow; the red plant used in the cross must therefore have been a heterozygote to give the observed 1:1 ratio in the F1. Selfing of the two plants used in the experiment was not succesful due to self-incompatiblility (which is very common in Iridaceae).
From this I concluded that yellow-flowered C. floribunda plants do not warrant formal taxonomic recognition. They are acyanic (i.e. anthocyanin lacking) variants, analogous to the white-flowered forms of many garden ornamentals that are no longer formally recognised except at the level of cultivars. This experiment was first published in December 1998 on my old website, which is archived here by the Wayback Machine, and the conclusion was also mentioned in the Horticultural Flora of South-Eastern Australia. My only rethink since then has been that there is a bit of variation in the intensity of the red flowers; possibly the more intense ones are homozygous for the wild-type allele and the paler ones with more yellow visible on the tube are the heterozygotes.
Bolus, L. (1933) Plants – new or noteworthy. South African Gardener 23: 46-47.
Cooke, D.A. (2005) Iridaceae. In Spencer, R.D., Horticultural Flora of South-Eastern Australia 5: 196-262.
This blog is a place where I’ll post occasional raves on subjects that interest me and may possibly attract another reader or two.
For the benefit of the search engines, I’m interested in plant taxonomy especially Iridaceae and Centrolepidaceae; breeding garden varieties of Watsonia; autistic culture; evolutionary theory that goes beyond genetic reductionism; Scientology; the politics of autism; Buddhism; and debunking junk science. Also (since I get paid for it) in applying botanical science to government policy on weed management as part of land management.
Some of my heroes and role models are: Lan CaiHe, ER Eddison, Padmasambhava, GK Chesterton, Agnes Arber, L Ron Hubbard, Vladimir Nabokov, HuiNeng, Ralph Vaughan Williams, Barbara McClintock, Olaf Stapledon, FaZang, Michael Polanyi, Aldous Huxley, Alfred Russel Wallace, John Coltrane, Emily Bronte, Simon Conway Morris, Pierre Bonnard, David Bellamy, Dennis Stephens, Michael Faraday, Lewis Mumford ….
I’m passionate about the need for our society to rediscover some intellectual roots. Mechanism and reductionism just won’t cut it anymore, but neither will a return to mysticism or superstition. By intellectual roots I mean knowing something real as a starting point for learning, and knowing how we know it: I’d call these things metaphysics and epistemology, if such words were not too oldfashioned for today’s hiphop-style academics.