Genetic Resources and Crop Evolution 51: 259–267, 2004.

259
 2004 Kluwer Academic Publishers. Printed in the Netherlands.

Phalaris canariensis is a domesticated form of P. brachystachys

R.N. Oram
CSIRO Plant Industry, GPO Box 1600, ACT, 2601 Canberra, Australia (e-mail:
rex.heather@ webone.com.au; phone: 161 2 6281 3654)

Received 5 April 2002; accepted in revised form 14 October 2002

Key words: Annual grass, Bird seed, Cereal, Dimorphic seeds, Domestication, Non-shedding mutant

Abstract

Three hybrid plants were obtained by hot-water emasculation of Phalaris canariensis L. panicles immediately
before anthesis, followed by the daily application of pollen from P. brachystachys Link. The hybrids were fully
fertile, and the seed retention characteristic of P. canariensis was inherited in the F 1 , F 2 and F 3 generations as a
single recessive trait, showing that P. canariensis is a cultigen derived from P. brachystachys. The dominant allele
for seed shedding is designated Ssh. All 1008 plants examined in segregating F 3 families had either short sterile
florets and shed their seeds, or long sterile florets and retained their seeds, i.e. there were no recombinants. No seed
retaining BC 1 F 2 plants could be found in the progeny of (P. truncata Guss.3P. canariensis)3P. truncata,
suggesting that the ssh ‘allele’ may be a number of mutant alleles in a segment of chromosome in which
recombination is suppressed in hybrids of P. canariensis with P. brachystachys, but in not those with P. truncata.
Compared with P. brachystachys, P. canariensis has heavier seeds, stronger panicle branches and thicker
peduncles. These traits were polygenically inherited in F 2 and thus appear to have been developed by the
accumulation of alleles during cultivation of P. canariensis as a grain crop. Modified seeds with much reduced
sterile florets were found in the terminal spikelets of most panicle branches of P. brachystachys. Unlike the typical
seeds, the modified seeds were often retained in the outer glumes when the panicle fragmented. It seems that this
system allows these seeds to be dispersed much further by animals, wind or water than typical seeds. This
mechanism has been lost in P. canariensis.

Abbreviations: n – gametic chromosome number, CPI – Commonwealth Plant Introduction

Introduction characteristic of P. canariensis permits its cultivation

Phalaris canariensis L. is the only member of a genus
containing 22 species in which the ripe seeds are
retained in the panicles after maturity (Baldini 1995).
In the other 13 Old World species, and in all of the
eight New World species, the mature fruits consisting
of a fertile floret and two reduced, sterile, basal
florets, which are termed ‘seeds’ in this paper, are
shed from the outer glumes soon after they mature.
An exception is a mutant, seed-retaining form that has
been found in the perennial pasture grass, P. aquatica
L. cv. Australian (McWilliam and Gibbon 1983). This
type of seed retention depends on homozygosity of
recessive alleles at four loci. The seed-retaining
as a grain crop, the grain being valued particularly for
feeding caged birds. Formerly, its flour was used as a
size for textiles, and, in some south-western European
countries, as a farinaceous food or blended with wheat
¨
flour for bread-making (Kornicke 1885). Febrel and
Carballido (1965) pointed out that P. canariensis is
still cultivated to a limited extent as a food grain in
Spain. de Wet (1979, 1992) argued teleologically that
P. canariensis was domesticated in south-western
Europe from a wild form of the same species, but at
those dates there was no direct evidence to support the
claim.
P. canariensis is one of three Phalaris species with
six chromosomes in the gametic set (Ambashta 1956).
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All of the other naturally occuring species have n 5 7,
14 or 21 (Baldini 1995). Two self-pollinating, annual,
n 5 6 species (P. canariensis and P. brachystachys
Link) resemble each other closely. Both have been
crossed with the third n 5 6 species, the taller,
perennial, largely self-incompatible species, P. trun-
cata Guss., but the F 1 s are partially sterile due to
incomplete chromosome pairing at meiosis (Cialzeta
1967a, 1967b). However, the two self-pollinating
species could not be hybridised (Cialzeta 1967a).
The principal morphological differences between P.
brachystachys and P. canariensis are the length and
morphology of the lemmas of the two sterile florets at
the base of the fertile floret, and in the retention or
shedding of the ripe seeds (Anderson 1961; Baldini
1995). In P. brachystachys, the sterile lemmas are
short, broad and have a fleshy appearance (Figure 1),
whereas those of P. canariensis are chaffy and about
one-third as long as the fertile lemma (Figure 1).
These marked and consistent differences have led
most taxonomists to place the taxa in different
species; only one of the 15 published descriptions
listed in the latest revision of the genus (Baldini 1995)
reduced one species, P. brachystachys, to varietal
status within the other.
This paper describes the hybridisation of P. brac-
hystachys and P. canariensis, and the segregation in
the F 2 and F 3 generations of the characters used to
distinguish them. This information clarified the rela-
tionship between the wild and domesticated taxa,

Figure 1. Mature spikelets of Phalaris brachystachys (a) and P. canariensis (b), showing the short, fleshy, basal sterile florets, the convex bases
of which (arrowed) pinch off the rachilla of the former species, and the long, chaffy sterile florets of the latter species, in which the concave
bases bend but reinforce the rachilla. The scanning electron micrographs below the light microscope photographs were provided by Prof. J.R.
McWilliam, and show the calluses (arrowed) at the bases of the sterile florets at higher magnification. The outer glumes of the P. canariensis
spikelet were cut off the pedicel to show its attachment to the rachilla. Magnifications: a (upper) 22 3, a (lower) 38 3, b (upper) 15 3, b
(lower) 26 3.

showing that they belong to the same biological
species and that the two sterile lemma length / seed
shedding traits are controlled by two different alleles
at a functionally single locus. The segregation of
some other morphological differences between the
two forms also was examined.
Materials and methods
Hot water emasculation
Two days before the beginning of anthesis, in mid-
morning, two panicles of the Moroccan P. canariensis
accession, Commonwealth Plant Introduction (CPI)
No. 14690, were plunged into distilled water at 46 8C
in a wide-mouthed Dewar flask. The panicles were
withdrawn after 5 min, when the water temperature
was 44 8C. Anthesis in the treated panicles was
delayed for several days, but after four days anthesis
appeared imminent. Each treated panicle then was
enclosed in a glassine bag with a flowering panicle of
P. brachystachys, CPI 14830 (Israel), which was
removed when anthesis was completed. Some P.
canariensis anthers then appeared to have shed pollen
while the panicles were bagged.
Observations on three generations of hybrids
Progeny from the only fertile treated panicle were
grown to maturity in a glasshouse and scored for seed
retention. The three plants which shed their seeds
were considered to be putative F 1 hybrids. Their self-
pollinated progeny were grown with representative
plants of the parental species. The F 2 plants were
allowed to self-pollinate and the panicles bagged
before seed shedding began. The traits measured on
the parental and 39 F 2 plants were: pollen stainability
by cotton blue in lactophenol, seed retention, stem
diameter 2 cm below the panicle, panicle length,
length and breadth of the fertile floret, length of the
two basal sterile florets, and seed weight. Measure-
ments on the smaller structures were made with an
ocular micrometer in a dissecting microscope. The
strength of a representative lower branch on the ovate
to sub-ovoid panicles was measured by clamping each
panicle at the base at 458 to the horizontal and
applying 20 and 50 g weights successively to the tip
of the branch until it broke. For statistical analyses,
the ratings of ,20, 20, 20–50, 50 and .50 g were
interpreted as 15, 20, 35, 50 and 60 g, respectively.
261
The pollen mother cells of a representative F 2 plant
were examined at meiosis. To examine the synchrony
of tillering, the date of first flowering on successive
tillers was recorded for 18 plants, four of them with
long sterile florets. An additional 43 F 2 plants were
grown, self-pollinated, and scored for sterile floret
length and seed retention. A set of 66 F 3 families was
chosen at random, and 60 seeds of each were sown in
single row plots in a field nursery. At maturity, each
plant was scored for sterile floret length and the level
of seed retention. The extent of seed retention in a
sample of 18 plants was determined by bagging a
panicle on each before shedding began, and, at ma-
turity, tapping the bagged panicle firmly against the
side of a collecting dish three times. The weights of
the shed and retained seed were measured.
The segregations observed in the F 2 and F 3 genera-
tions were tested for goodness of fit to expected ratios
by x 2 analyses, using Yates’ correction factor for
two-class segregations totalling less than 200 plants
(Little and Hills 1978).
Exceptional seeds in P. brachystachys
Seeds with atypically short sterile florets and sparse
hairs on the fertile lemmas were observed in P. brac-
hystachys samples. Panicles were dissected to de-
termine the distribution of these seeds and the charac-
teristics of their subtending glumes. A search was
made for the corresponding modifications in P.
canariensis.
Results
F1 hybrids
Very few anthers were exserted from one of the
panicles treated with hot water, and no seeds were
produced. Some upper florets in the other panicle
remained closed, but the remainder exserted anthers
and stigmas, were pollinated and most set seeds. Of
the 25 plants grown from these seeds, 22 retained
their ripe seeds like the female parent and thus re-
sulted from self-pollination. But three offspring shed
their seeds, suggesting they were outcrosses which
received a dominant gene for seed-shedding from the
P. brachystachys male parent. Also, these seeds re-
sembled those of the male parent in having short,
fleshy sterile florets.
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F2 generation

The pollen of 39 F 2 plants contained 0–16% empty or

partly filled grains (Table 1). Thirty-five of these
plants had short sterile florets like P. brachystachys
seeds, whereas 4 plants had long sterile florets like P.
canariensis. In the whole F 2 population, the numbers
of plants with short or long sterile florets were 67 and
15, respectively, a segregation which fits a 3:1 ratio
(x 2 5 1.63, P 5 0.220.3). The heterogeneity x 2 for
the segregations in the three families was 3.08, a
non-significant value (P 5 0.220.3). All plants which
retained their seed had long sterile lemmas on the
basal florets and all plants which shed their seeds had
short sterile florets. Figure 2 shows that there was
some variation in the length of the lower sterile florets
but no overlap between groups with long and short
sterile florets. The mean lengths of the upper and
lower sterile florets in the short-floretted set were 1.05
and 0.93 mm, respectively, and these values closely
resembled those for P. brachystachys (1.08 and 0.97 Figure 2. Joint distributions of the mean weight per seed and length
mm, respectively). The corresponding means in the of the lower sterile floret for P. canariensis (j), F 2 plants with long
sterile florets (h), P. brachystachys (d), and F 2 plants with short
set with long sterile florets were 2.60 and 2.90 mm,
sterile florets that were either homozygous (쏻) or heterozygous (쏍)
respectively, whereas the values for P. canariensis for the dominant wild type allele, Ssh.
were 2.45 and 2.64 mm. The lengths and widths of the
fertile florets and the weight of the spikelets con-
taining a seed in the two F 2 sterile floret length classes florets were no more synchronous for the beginning of
were intermediate between, but tended to resemble flowering on successive tillers than the short-floretted
the respective parental values (Table 1). There was plants.
considerable overlap between the values for the two Meiosis was normal in the only F 2 plant examined.
classes, as illustrated for weight per seed in Figure 2. At diplotene and diakinesis, all chromosomes were
However, of these traits, the only one which differed paired in ring or rod bivalents. Only one nucleolus
significantly between the classes was weight per seed. was formed. Of the 150 anaphase II and telophase II
The distributions of outer glume length and tiller cells observed, only two contained a laggard chromo-
number per plant of the two seed retention / sterile some. One laggard appeared to be an entire chromo-
floret length phenotypic classes overlapped extensive- some and the other an acentric fragment. Micronuclei
ly. Panicle branch strength means differed signifi- were not observed at the first pollen grain mitosis.
cantly between the two phenotypes and were corre- These observations, together with those on the high
lated with the respective parent values, but negative fertility of the F 2 and F 3 plants, show that the
correlations were shown in the case of peduncle genomes of P. canariensis and P. brachystachys are
diameter and date of anthesis (Table 1). virtually identical.
The synchrony of anthesis on successive tillers was
much higher on P. canariensis plants than on those of F3 generation
P. brachystachys - the former produced only a few
tillers, whereas the latter produced tillers over a long All 250 of the F 3 plants derived from 12 long-floretted
period. The F 2 plants which began flowering in late F 2 plants were themselves long-floretted. However,
August produced new tillers that began flowering 32 of the F 3 families from short-floretted F 2 plants
until early or mid November, whereas plants begin- segregated for sterile floret length, and 22 F 3 families
ning flowering in late October or early November bred true for short florets. The frequencies of the three
developed newly-flowering tillers only over intervals kinds of F 3 families fitted a 1:2:1 segregation ratio (x 2
of 7–10 days. The four F 2 plants with long sterile 5 3.09, P 5 0.220.3), confirming that the major
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Table 1. Seed, panicle and plant characteristics differentiating Phalaris brachystachys (P. brach) and P. canariensis (P. can), and their
associations with sterile floret length in F 2 .
Character Parents F 2 sterile floret class
P. brach P. can Short Long
Range Mean Range Mean
Pollen stainability (%) 98.0 91.0 84–100 94.9 95–100 96.7
Fertile floret length (mm) 4.74 5.25 4.26–5.25 4.67 4.68–5.33 5.08
Fertile floret width (mm) 1.55 2.44 1.56–2.03 1.81 1.69–2.50 2.02
Weight per seed (mg) 3.58 9.40 3.9–7.3 5.21 4.6–8.8 6.30*
Outer glume length (mm) 7.27 8.50 6.9–8.9 7.93 7.0–8.0 7.63
Panicle branch breaking force (g) 15.8 60.0 15–60 37.6 35–60 53.8*
Peduncle diameter (mm) 0.59 0.81 0.57–0.87 0.70** 0.52–0.65 0.59
Date of anthesis (December) 20.3 10.0 26–30 10.4 12–30 21.0*
Tiller number 15.2 4.0 8–19 10.8 6–17 11.7
*,** indicate the mean which is significantly greater, at the 5% and 1% levels of probability, respectively, than the mean in the other floret
length class, as determined by t-tests

taxonomic difference between the parental species is
controlled by two alleles at one locus. The allele for
short sterile florets and seed shedding in the wild
parent is dominant to the allele in P. canariensis. The
dominant allele for seed shedding is designated Ssh,
and the recessive, ssh. The 1:2:1 segregations in the
descendants of the three F 1 plants were not signifi-
cantly heterogeneous (x 2 5 2.54, P 5 0.520.7). Of
the 1008 plants classified in the segregating F 3
families, 758 had short sterile florets and 250 had long
ones, a very close fit to the expected 3:1 ratio (x 2 5
0.02, P 5 0.820.9). The segregations in the family
groups descended from the original three F 1 plants
were homogeneous (heterogeneity x 2 5 0.93, P 5
0.520.7). The proportion of seed retained by a sample
of 15 long-floretted plants ranged between 90 and
99.5%, except for one plant with 84.4% retention. In
this plant the concave calluses on the base of the
sterile florets flattened the rachillae and bent them at a
sharp angle, making the rachillae more likely to break
when the panicle was jolted. Most of the eight, short-
floretted plants examined shed more than 50% of their
seed, the maximum retention being 63.5%. In this
plant, the convex calluses on the lower sterile florets
failed to grow completely up to the upper calluses,
and hence the rachillae were not always completely
pinched off.
Modified spikelets
The mature fruits of CPI 14830 P. brachystachys were
found to be dimorphic. Most had the morphology
described by taxonomists, which is illustrated in the
left and centre images in Figure 3, but 5.0–9.8% of
the seeds had shorter, less hairy flowering glumes and
very much reduced sterile florets, as shown on the
right in Figure 3. The lemmas of the sterile florets
were reduced to minute scales, and the calluses were
reduced to about half the usual size. These modified
seeds frequently were brown in colour, whereas the
typical seeds were usually a darker, grey colour. The
seeds produced by plants grown from the two types of
seeds were similarly dimorphic.
A modified seed was only found in a sessile or
sub-sessile terminal spikelet in a group of five, seven
or nine spikelets, provided the terminal spikelet was
flanked by a pair of unequally pedicellate spikelets, as
shown in Figure 4. The remaining one, two or three
pairs of spikelets in the group each consisted of one
almost sessile and one pedicellate spikelet. The more
sessile member of each set of two or three spikelets
was distal and abaxial to the others in the set (Figure
4). The less well developed panicle branches near the
bottom and top of the panicle did not contain modified
seeds. Modified seeds also were found at frequencies
of 3 and 12.5% in samples from two F 2 plants with
short sterile lemmas. Some CPI 14690 P. canariensis
seeds also appeared to be modified in flowering glume
length and hairiness, but the differences were too
small to accurately classify all seeds. No modifed
seeds could be found in the panicles of a plant grown
from commercial bird seed, which probably was
produced by an improved cultivar of P. canariensis in
Queensland, Australia. All of the P. brachystachys
spikelets in a group, except the terminal one, had a
strip of red-brown tissue on the inner surface at the
base of each of the outer glumes along the junction
with the pedicel. A groove occurred at the same
position on the outer surface (Figure 4). At maturity,
the coloured strip and groove appeared to act as a
264

Figure 3. Typical (left and centre) and atypical (right) seeds of P. brachystachys. The two sterile florets have been removed from the spikelet at
the left. The rachillae of both seeds have been pinched off by the calluses at the base of the sterile florets. The atypical seed has much reduced
sterile florets, and the fertile floret has a shorter, less hairy lemma and palea. Its rachilla was not pinched off by the sterile florets, but fractured a
short distance below them, leaving a stump (arrowed).

hinge about which the outer glumes opened, perhaps
initially under pressure from the bent sterile lemmas
(Figure 1), allowing the seed to fall from the spikelet.
If the terminal spikelet produced a modified seed, it
lacked the coloured ‘hinge’ tissue in the outer glumes,
and, although the glumes had outer grooves at the
base, they remained closed and the seed usually was
retained within them.
Discussion
Hybridisation of P. brachystachys and P. canariesis
has shown that these two taxa belong to a single
biological species, P. brachystachys being the wild
ancestor and P. canariensis the cultivated form de-
rived from it by a single dominant to recessive muta-
tion that resulted in the suppression of the shedding of
ripe seeds from the panicles. This genetic change is
analogous to those preventing seed shedding during
the domestication of the major temperate cereals
(Zohary 1973).
Several other morphological and physiological
changes appear to have been made subsequent to the
initial cultivation of the mutant form, particularly a
reduction in tiller number, an increase in synchrony of
tiller production, an increase in seed size and the
suppression of the wild system for the production of
modified seeds, which are less readily shed from the
outer glumes. Apart from the latter trait, the dis-
tributions of these traits in F 2 were unimodal, e.g.,
weight per seed in Figure 2, indicating that each was
controlled by alleles with small effects at a consider-
able number of loci. These changes parallel those in
the other domesticated cereals, which were reviewed
by de Wet (1979). Although P. canariensis is a minor
crop, genetic improvement of it continues in some
regions. Thus, Robinson (1983) reported the release
of a new cultivar with higher grain yield and bulk
density in Minnesota. An induced mutant with very
few hairs on the fertile lemma and palea has been
commercialised in Canada as cv. CDC Maria to avoid
irritation to grain producers during harvest and the
need to remove the hairs mechanically for use as bird
seed (Hucl et al. 2001). The hairs of several annual
phalaris species, including P. canariensis, contain
silica fibres with fine points of similar size and shape
as those of asbestos fibres. These siliceous fibres have
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Figure 4. Seven spikelets from one mature panicle branch of CPI 14830 P. brachystachys viewed from the abaxial side. The terminal spikelet is
almost sessile, has shorter outer glumes which remain almost completely closed, and contains a modified seed (arrowed), as shown in Figure 3.
The other three pairs of spikelets are unequally pedicellate, the outer glumes have opened by lateral rotation at the sutures at the junction of the
outer glumes and pedicels, and have shed their seeds. In each set of two or three spikelets, the one with the shorter or shortest pedicel is adaxial.

been implicated in the high incidence of oesophageal
cancer in north-eastern Iran, where they have been
found as a contaminant in bread (Sangster et al.
1983).
Although only one accession each of P. brac-
hystachys and P. canariensis have been hybridised,
these were typical members of the taxa from their
Mediterranean homelands, so crosses between other
pairs of accessions probably would reveal the same
genetic differences. However, crosses within and
between the taxa were very difficult to make. Al-
though emasculation with forceps appeared to leave
the glumes and pistils undamaged, cross pollination
invariably gave no seed. Similarly, pollination of the
stigma tips, which emerge from the flowering glumes
several hours before the anthers, yielded no hybrids.
Two additional attempts to hybridise the species were
made, using hot water emasculation, in order to
examine meiosis and gamete fertility in F 1 hybrids,
but in both cases no seeds were set, apparently be-
cause the heat treatment used previously was too
damaging on the later occasions.
The ssh allele for seed retention has manifold
effects on the development and final morphology of
the sterile florets, particularly the trebling of the
length of the lemmas and the change in shape of the
basal calluses from convex pincers, which constrict
and break the rachilla, to concave structures which
grow alongside the rachilla and strengthen it. Al-
though over a thousand plants were scored for seed
retention and sterile floret length in segregating F 3
families, no seed-shedding, long-floretted or seed-re-
taining, short-floretted recombinants were found. This
shows that ssh is a mutation at a single pleiotropic
locus, perhaps with a regulatory effect on several
other loci, or a deletion of a segment containing two
or more loci, or a multi-locus segment in which
recombination is suppressed in both P. brachystachys
and its derivative. Evidence for the latter possibility
was obtained from backcrosses of P. canariensis to P.
truncata Guss. that were made to transfer the seed
retention trait to the perennial species, which is a
useful forage plant on alkaline, clay soils in parts of
the Mediterranean region (Le Houerou ´ 1979). Al-
though a large population was examined, no com-
pletely seed-retaining plant was found in the BC 1 F 2
266
generation. Partial retainers were selected and inter-
crossed in three successive generations, but complete
retainers were not found in the progeny, suggesting
that ssh is a segment containing many genes with
different effects which recombine freely, or ssh reg-
ulates other genes differently, in P. truncata3P.
canariensis derivatives. To have been selected by
ancient cultivators, the original mutant must have
been fully seed-retaining, or almost so. Whatever the
nature of the Ssh and ssh ‘alleles’, their effects on the
structure of the calluses on the sterile florets varied
slightly in the F 3 generation: one Ssh / - plant retained
more seed than the other 14 plants examined due to
incomplete closure of the calluses, and one out of
eight ssh /ssh plants shed more seed than usual be-
cause the calluses constricted and bent the rachillae to
an unusual extent. This variation may depend on the
joint action of several mutant alleles in the back-
ground genotype.
Most, if not all, of the other differences between P.
brachystachys and its seed retaining derivative appear
to have been generated by fixation of variant alleles
after the domesticating mutation occurred. None of
the traits examined was closely correlated with those
directly controlled by ssh, but rather they showed
multigenic inheritance in F 2 . The modified seeds from
the terminal florets of many P. brachystachys panicle
branches may also follow this pattern - although some
seeds of CPI 14690 P. canariensis had some short-
ening of the sterile lemmas and reduction of hair
number on the fertile glumes, the changes were much
less distinct than in P. brachystachys, and could not be
observed in the seed of a modern commercial cultivar.
It seems that the modification has selective value in P.
brachystachys, but not in the seed retaining P.
canariensis. The value of the modified seed to a wild
species probably depends on its much more frequent
retention in the spikelet than the other seeds in the
panicle branch. The retention of a seed in the terminal
triplet of spikelets may permit its dispersal over a
much longer distance than is possible for the other
seeds, perhaps in the wool or hair of grazing animals,
or on currents of air or water. By contrast, the ssh
mutant mainly has been transported by man, either
deliberately for re-sowing at favourable sites, or ac-
cidentally in refuse or ships’ ballast (Anderson 1961).
The function of the brown band on the inner surface at
the base of the outer glumes of florets which shed
their seeds, but not in the florets which usually retain
their seeds, remains unknown, as does the role of the
fleshy sterile lemmas which occur only on the shed
seeds. However, these correlations suggest that the
fleshy lemmas arch outwards as they dehydrate (see
Figure 1) and help to force the outer glumes apart.
These glumes appear to rotate about the coloured
‘hinges’ at their bases. Bonin and Goplen (1963)
observed constrictions similar to the grooves on the
outer glumes of P. brachystachys at the bases of some
of the outer glumes of a clone of P. arundinacea L.
that shed 44% of its seed; these constrictions were
absent in a clone that shed only 19% of its seed.
There appears to be limited variability within the
genepool of P. canariensis. For example, the cultivars
‘Super Mammoth Spanish’, ‘Lyons’ Selection’ and
‘Nunbank’ from Queensland were very similar in seed
and head lengths and widths, weights per seed and
peduncle diameter when grown in Canberra (Oram,
unpub.). Also the samples from six Spanish wine-
growing provinces examined by Febrel and Carbal-
lido (1965) were very similar in weight per seed, and
in oil, protein and crude fibre concentrations. Similar
conclusions were drawn from the analysis of variants
in eight enzymes in 40 accessions of P. canariensis,
24 accessions of P. brachystachys, and 70 accessions
of three other annual Phalaris species (Matus and
Hucl 1999). As is common in domesticates, less
polymorphism was found among accessions of P.
canariensis than in P. brachystachys and the other
wild and weed species. Furthermore, the P. canarien-
sis accessions were embedded within those of P. brac-
hystachys in the phenogram of Jaccard’s similarity
coefficients (Matus and Hucl 1999), strongly support-
ing the evidence reported here for the origin of P.
canariensis as a mutant within P. brachystachys.
The low level of variability among improved cul-
tivars and volunteer populations of P. canariensis
reduces the opportunities for further genetic improve-
ment, and the difficulty of crossing different geno-
types compounds the problem. Additional studies on
crossing techniques are needed. A more gentle meth-
od of emasculation which may be effective is that
developed by Knight (1966) for Dactylis glomerata
L. If no satisfactory emasculation method can be
found, a single recessive, male-sterile mutant could be
induced and used as the female parent in a wide range
of crosses, followed by a recurrent selection program
using male sterile segregates as female parents in each
crossing generation (Gilmore 1964). Some introgres-
sion of genes from diverse P. brachystachys acces-
sions also would be required to broaden the gene pool.
Collectively, these methods should accelerate the
continuous genetic improvement of P. canariensis.
 267

Acknowledgements ¨
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