Both the typical yellow form of Freesia leichtlinii (Dune Freesia) and the white form (previously known

as Freesia alba) are important in the origin of modern freesia cultivars.

Scentsational freesias

Chemical analysis of the fragrance of freesias supports the division of the genus Freesia into two groups
based on DNA and bract texture
by John Manning and Roman Kaiser

or city dwellers around the world, the appearance of freesias in

florists shops and in the barrows of flower sellers on street corners
is the urban equivalent of the unfurling of spring wildflowers
in fields and woodlands. Unlike many of their rural cousins, however,
freesias pour forth a strong, sweet fragrance from their trumpet-shaped
flowers, sounding a brilliant olfactory fanfare to dispel the dreary
northern winter. Their bright sprays of blossom burst onto streets with
the lan of a cancan chorus line, the ruffled flowers dancing down each
stem in a perfectly choreographed row, petals swirling around dainty,
creamy stamens vivacious and irresistible as the grisettes of Maxims.
Modern freesia cultivars, puffed up by polyploidy, are essentially the
product of the ingenuity and application of late-Victorian and Edwardian
breeders in Europe. The wild species from which their stocks were
originally drawn are endemic to the south-western Cape but are barely
known, even in their native land.
Botanists today recognize 16 species of Freesia, one of them so
recently recognized that it has yet to be formally named. All but four of
the known wild species are endemic to Namaqualand and the southwestern Cape, with the greatest concentration in the Worcester-Little
Karoo and adjacent littoral, in the Overberg and the southern Cape.
Known colloquially as flissies from their flask-shaped flowers arranged
in perky, arched spikes, most species exude a heady, violet-like fragrance
during the day.

132

VELD&FLORA | SEPTEMBER 2009

Tentative beginnings
The first freesias found their way to Europe in the mid Seventeenth
Century at a time of rising interest in Cape plants. Bulbs and succulents
were among the plants that most readily survived the rigours of the
long sea voyages from the Cape to the ports of Europe, and the first
two Freesia species known to science were described from plants that
had been established in cultivation in The Netherlands some time
prior to 1768. They were given names that alluded to some notable
characteristic, aptly in one case the striking fragrance for which the
genus would later become famous. In naming F. caryophyllacea, the
Carnation-scented Freesia, however, the Dutch botanist Nicolaas Burman
reveals a deplorable knowledge of bouquet since no species of Freesia
produces oil of cloves, the primary component of the smell of wild
carnations. In keeping with botanical conventions of his day, Burman
placed his two new species in the genus Gladiolus on account of their
two-lipped, trumpet-shaped flowers.
Freesia itself was only established as a separate genus a century later,
in 1866, by the German botanist, F.W. Klatt, who is honoured in another
botanical curiosity from the Cape, the shrubby iris Klattia. At this time the
name Freesia was applied only to those species with the characteristic
funnel-shaped flowers that are familiar to us from the cultivated forms,
but three decades later Klatt expanded his conception of the genus to
include a few species with narrowly tubular, mostly unscented flowers,

notably the Forest Painted-petals, F. grandiflora and F. laxa, both with

brilliant scarlet, salver-shaped blooms. It is this circumscription that we
follow today.
By the end of the nineteenth century, just over half of the currently
accepted species of Freesia were known to science, all of them collected
and described by European botanists. The establishment of a botany
department at the South African College (later the University of Cape
Town) in 1903 and of Kirstenbosch Gardens (now Kirstenbosch National
Botanical Garden) ten years later, marked the beginning of indigenous
scientific study in South Africa, leading directly to the discovery and
description of four more species of Freesia in the first decades of the
Twentieth Century. Here the total remained until the opening years
of this century, during which another three highly localized species
will have been named, bringing the number of species of Freesia to its
current total. These 16 names are not, however, the only ones to appear
in the botanical and horticultural literature as many Freesia species were
known under more than one name to different botanists.
Early botanical accounts of Freesia bear little resemblance to our
current understanding of the genus, and this confused legacy is
especially pernicious in attempts at understanding the early history of
the cultivars. The first attempt at a comprehensive treatment of Freesia
in its classic sense was that of the late-Victorian British botanist, J.G.
Baker, who recognized just the single species Freesia refracta, which he
subdivided into three varieties that we recognize today as three distinct
species. Bakers understanding of the genus was derived primarily from
information supplied by the amateur horticulturist, M. Foster, Professor
of Physiology at Cambridge, but despite their best efforts, the exact
application of some of the other names current at this time remained
uncertain. The misuse of the name F. refracta in the horticultural literature
for forms of F. corymbosa and F. leichtlnii stems from this time true
F. refracta was never used in Freesia breeding. The naming of several
new species of Freesia in the early decades of the Twentieth Century

by the South African botanist H.M.L. (Louisa) Bolus stimulated the

first modern treatment of Freesia by the English botanist, N.E. Brown,
who distinguished no less than nineteen species, many of them now
understood to be nothing more than minor colour and leaf variants.
Brown was, however, the first to recognize the important difference in
the texture of the floral bracts that has turned out to be so pivotal in our
modern classification of the genus. Browns plethora of poorly defined
species was consolidated in the last decades of the Twentieth Century by
Peter Goldblatt, who also succeeded in pinning down several of the early
names that had confused earlier botanists.
The modern synthesis
By now it was accepted that freesias came in two very different floral
forms but it was not clear exactly how these two types related to one
another. The first attempt at an evolutionary analysis focused on floral
characteristics, and it confirmed the prevailing idea that the classic,
funnel-flowered species comprised a discrete lineage of closely related
species. This contrasted sharply with the situation among the tubularflowered species, which fell into three different groups, indicating a far less
intimate relationship among them. These results suggested that earlier
botanical practice, which grouped the four tubular-flowered species
together in the genus Anomatheca, required serious reconsideration.
The development of DNA-based techniques for studying relationships
among organisms offered a chance to resolve these two conflicting
ideas. Carried out in conjunction with Dr Flix Forest at Kew, the picture
that emerges is fascinating and provocative.
Like the morphological analysis, the DNA analysis places the tubularflowered species in three separate lineages but rather than forming a
loose alliance set apart from the trumpet-flowered species, the tubularflowered plants appear to be closely entangled among the classic freesia
flowered species. In fact, the primary division among species of Freesia
coincides not with flower shape at all but exactly with the difference in

LEFT: Freesia speciosa, a rare species from the western Little Karoo, has the largest flowers of all wild freesias. RIGHT: Freesia grandiflora, a widespread species of tropical and subtropical woodland, with
unscented, scarlet flowers, was previously placed in the genus Anomatheca on account of its slender floral tube. Photos: J. Manning.
SEPTEMBER 2009

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133

Freesia leichtlinii, Kirstenbosch, 28.7.05, 9.00-15.00

-Terpineol

35
30
25
20
15
10

OH

70
65
60
55
50
45
40
35
30
25
20
15
10
5

26

28

30

32

34

36

38

40

42

44

46

48

50

Time (min)

K63055
30m CP-Wax 52CB, 2 l on column
100

OH

50
45
40
35
30
25
20

OH

OH

OH

OH

OH

80
75
70
65
60
55
50
45
40
35
30
25

16

18

20

22

24

26

28

Time (min)

30

32

34

36

38

40

42

44

46

50

Freesia x hybrida white, 18.7.06, 9.40-13.40

16

18

NL:1.57E9
m/z= 33-350

15

10

10

10

12

14

16

18

20

22

24

26

28

30

32

34

36

38

40

42

44

46

48

50

Time (min)

10

12

14

20

22

24

26

28

Time (min)

30

32

34

36

38

40

42

44

46

Gas chromatograms of selected Freesia species and cultivars. Individual peaks represent different chemical compounds, and the size of each peak is proportional to the relative concentration of that
chemical.
A The scent of F. leichtlinii is characterized by the presence, among other compounds, of high concentrations of -terpineol and -ionone as well as smaller amounts of three other diagnostic
chemicals (Dihydro--ionone, 2-methylbutyraldoxime (E+Z) and -selinene), all highlighted in red. These particular compounds are absent from the scent of F. corymbosa, which is recognizable by
the presence of trace amounts of four unusual chemicals derived from the more common, dominant constituent, linalool (2-methyl-2-vinyltetrahydrofuran-5-ol, 2,6-dimethylocta-3,7-dien-2,6-diol
and 2,6-dimethylocta-1,7-dien-3,6-diol), all highlighted in blue.
B White-flowered commercial Freesia cultivars betray their mixed parentage by combining the diagnostic fragrance markers of both F. leichtlini (highlighted in red) and especially F. corymbosa
(highlighted in blue), whereas yellow-flowered cultivars approach pure Freesia leichtlinii in their fragrance.
the texture of the floral bracts that had first been pointed out by Brown
in 1935. On the one hand we have those species with firm-textured,
green bracts, and on the other those with thinner, more membranous or
even papery bracts. Both groups are dominated by species with funnelshaped flowers but each also contains one or more tubular-flowered
species. This coincidence between DNA and bract type is reflected in our
decision to recognize these two groups as subgenera.
A cultivated past
The history of modern freesia cultivars goes back to the late Nineteenth
Century, with the introduction into cultivation of yellow- and whiteflowered forms of F. leichtlinii (the latter under the name F. alba).
Selection from this material gave a range of white and yellow forms
that enjoyed some popularity but it was the introduction of pink- and
deep yellow-flowered forms of F. corymbosa (known respectively as F.
armstrongii and F. aurea) in the closing years of the century that provided
the real stimulus to freesia breeding. By the end of the first decade of
the Twentieth Century the Dutch nursery firm of Van Tubergen had
produced a range of tall freesias in a variety of colours including blue,
mauve, shades of rose, yellow and white, and the first polyploid, Freesia
Buttercup, made its appearance in 1911. Today most registered cultivars
are polyploid, with diploid strains important mainly in areas where plants
are grown every season from seed.
Breeding of Freesia continues today with the resultant varieties,
including double-flowered types, bearing less and less resemblance to
the wild species from which their stock originally stemmed. The cultivars
range from 1030 cm in height, in habit from neat and tufted to tall and
graceful, in inflorescence from short and semi-erect to long, sparsely
branched and horizontal, and producing scarcely to strongly fragrant
flowers in almost every conceivable colour. Almost shockingly, however,
no additional genotypes appear to have been introduced into the mix
since those first crosses were made in the late Nineteenth Century, and
all commercial freesias are essentially derived from repeated crossings of
134

48

20

15

14

95

Relative Abundance

55

12

85

2,6-Dimethylocta-1,7-dien-3,6-diol

60

OH

2,6-Dimethylocta-3,7-dien-2,6-diol

65

cis-Linalool oxide (furanoid)

70

trans-Linalool oxide (furanoid)

75

Tridecan-2-one

80

10

90

2-Methyl-2-vinyltetrahydrofuran-5-one

85

2-Methyl-2-vinyltetrahydrofuran-5-ol

Linalool

90

100

NL: 1.37E9
m/z= 33-350

OH

95

K7704
30m CP-Wax 52CB, 1ul on column
RT: 2.30 - 50.00

Freesia corymbosa "armstrongii", 27.9.07, 11.00-16.00

RT: 2.30 - 50.00

2,6-Dimethylocta-1,7-dien-3,6-diol

24

2,6-Dimethylocta-3,7-dien-2,6-diol

22

-Ionone

20

-Terpineol

18

2-Methyl-2-vinyltetrahydrofuran-5-ol

16

Linalool

14

cis-Linalool oxide (furanoid)

12

2-Methylbutyraldoxime

10

trans-Linalool oxide (furanoid)

(E)-Ocimene

Eucalyptol

Limonene

Relative Abundance

Linalool

75

-Ionone

40

80

Dihydro- -ionone

45

Eucalyptol

50

NL: 7.27E8
m/z= 33-350

-Terpineol
-Selinene

OH

85

Caryophyllene

55

Freesia x hybrida yellow,18.4.09, 11.00-14.00

100

2-Methylbutyraldoxime

60

K69618
30m CP-Wax 52CB, 1.2l on column
RT: 2.30 - 50.00
90

-Selinene

65

B.

95

OH

Linalool

70

Caryophyllene

75

Limonene

Relative Abundance

80

2-Methylbutyraldoxime (E+Z)

85

NL: 1.14E9
m/z= 33-350

(E)-Ocimene

(E)-Ocimene

90

-Ionone

95

Eucalyptol

100

Relative Abundance

RT: 2.30 - 50.00

Limonene

K52936
30m CP-Wax 52CB,1l on column

Dihydro- -ionone

A.

VELD&FLORA | SEPTEMBER 2009

colour forms of just two wild species, F. corymbosa and F. leichtlinii. This is
an almost incredible neglect of the potential that exists among the wild
species for the addition of new flower shapes and other characteristics.
Although freesias are famous for their fragrant flowers, typically but
inadequately described in horticultural literature as sweet-smelling, very
few cultivars have been developed primarily for their fragrance. Indeed,
many commercial varieties have lost their scent during the breeding
and selection processes, a cause for special concern among Chinese and
Japanese growers. The favoured colours by far in both Holland and Japan
are yellow, followed by white, and it is probably no coincidence that
these colours are typically the most fragrant.
Scenting success
The division of Freesia into two groups based on DNA and bract texture
has received spectacular support from our chemical analysis of floral
fragrance in the genus. The scent in freesia flowers emanates primarily
from the bright yellow or orange patches that decorate the lower lip of
the flowers like smears of turmeric. Scented members of the group with
delicate bracts, such as F. corymbosa, produce floral fragrances dominated
either by the chemicals linalool (which has an attractive floral-woody
smell) or nerol (fresh and rose-like with citrus undertones). Members of
the group with leathery bracts, which includes F. caryophyllacea and F.
leichtlini, are chemically more diverse, with scents that are dominated
by nerol or -terpineol (sweet-floral with pine-needle undertones) but
with the signal addition of the chemicals -ionone and dihydro--ionone
(green-woody, fruity and very floral with a distinct freesia smell). The
ionones, which are extremely potent with very low odour thresholds to
the human nose, have found an enormous appreciation in perfumery
during the past three decades and many of the famous fragrances
contain high amounts of them, often in ratios similar to those found in
nature. Relatively widespread among flowering plants, ionones have the
curious characteristic that they are invisible to around 10% of humans
who are unable to detect their odour through a genetic peculiarity. As

48

50

a result, people who cannot smell ionones will

perceive the scent of flowers containing these
chemicals in their fragrance spectrum quite
differently from those fortunate enough to
have active ionone receptors.
A new beginning
Our identification of ionone-negative and
ionone-positive groups in the genus raises
the exciting possibly of developing distinct
fragrance-lineages in Freesia. The very early
crossing of F. leichtlinii with F. corymbosa
effectively removed this potential from current
breeding programmes by unwittingly blending
the two fragrance-groups.
The new understanding of relationships
among the species that was provided by
DNA analysis opens up the opportunity for
selective breeding within each of the two
fragrance-groups to derive fragrance-cultivars
that can only enhance our enjoyment of this
wonderful genus.
It is most unfortunate, for instance, that true
F. refracta has never figured in the parentage
of commercial cultivars as it has a rich, roselike scent stemming from the chemicals nerol,
geraniol and citronellol. These rosy-floral and
citrus-related scent constituents harmonize
exquisitely with the fragrance of the existing
cultivars. Another great mystery is why F.
caryophyllacea, an attractive and relatively
common species that was among the very first
to be grown in Europe, has never been involved
in breeding programmes. It offers ravishing
rosy-floral fragrance notes via geraniol and
nerol and their derivatives.
Floral scents, elusive and evocative, add a
final touch of glamour to the beauty of many
of our wildflowers but they are a sadly underappreciated feature that richly rewards closer
attention.

The authors
Dr John Manning is a Senior Specialist Scientist at the
Compton Herbarium, South African National Biodiversity
Institute, Kirstenbosch. Dr Roman Kaiser is a chemist
at the Givaudan fragrances and flavours company in
Switzerland.
WHAT DOES THAT MEAN?
Diploid Two of each type of
chromosome
Genotype The genetic make-up of an
organism as opposed to its physical
characteristics
Littoral Of or on the seashore
Morphological Shape and form or
physical characteristics of an organism
Polyploidy Having more than the
usual number of chromosomes

TOP: Freesia fucata, a rare species from near Villiersdorp, has the highly fragrant, funnel-shaped flowers that are characteristic of the
genus. The scent in freesia flowers emanates primarily from the bright yellow patches that decorate the lower lip of the flowers like
smears of turmeric.
ABOVE: True Freesia refracta was never used in the breeding of freesia cultivars and references to it in the early horticultural literature
actually apply to F. corymbosa and F. leichtlinii. Photos: J. Manning.
SEPTEMBER 2009

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