the calculated dominance deviation val- The use of this short culm population what factors cause the
what factors cause the variation3610 and
ues. Epistatic deviations were tested ac-
cording to the procedure outlined by
Mather and Jinks,2 and this test was ap-
plied only to the combined test results.
Results
To determine cross differences, we made
comparisons between the same genera-
tions (except P,) (Table 1). Only the means
of the B, generation (82 cm in Cl com-
pared with 73 cm in C2) were significantly
different at the .05 probability level. We
did not detect statistical differences be-
tween the variances of the same genera-
tion in each cross. Therefore, differences
in the effect of the long parents (401, FB),
which had identical means (143 cm) and
similar variances, were minimal.
From the results given in Table 1 and
shown graphically in Figure 1, the short
culm trait was determined to be partially
dominant (/?/dvalues: Cl = 0.59, C2 = 0.69,
combined crosses = 0.64) and to be con-
trolled by three loci or effective genetic
factors (Cl = 3.0, C2 = 3.1, combined
crosses = 3.1). Moderate broad sense (Cl
= 0.57, C2 = 0.61, combined crosses =
0.59) and narrow sense (Cl = 0.50, C2 =
0.54, combined crosses = 0.52) heritability
values were found. Highly significant (x2
= 96.98, df = 3, P < .01) deviations re-
sulting from epistasis were found.
Discussion
At least six loci, which are described as
short straw, compactum, or dwarf and
could conceivably influence culm length,
have been reported in rye.3 Detailed ge-
netic information (number of alleles, dom-
inant relationships, linkage, etc.) on these
loci is extremely limited and imprecise.
Also, the assigned gene symbols are in-
consistent, and the verbal descriptions of
their specific effects are vague. In both
short straw and compactum loci (c/7, ct2),
short straw is recessive. Among the four
dwarf loci, dwarfing is recessive at two loci
(dw2, dw3) and dominant at the other two
loci (Dwl, Dw2).
Our results indicated that the short culm
trait was partially dominant, was con-
trolled by three loci showing epistasis, and
had moderate heritability. The genetic re-
lationship between the six loci mentioned
above and the short culm population we
used is unknown. With the difference in
dominance relationships, epistasis might
be expected when some of these genetic
factors are combined.
5 0 6 The Journal of Heredity 1989:80(6)
as a source for developing commercial
short culm cultivars is being considered.
With this degree of genetic complexity and
the self-incompatibility present in rye, the
incorporation of this desirable character
would be difficult. However, the moderate
heritability values suggest that environ-
ment would be a relatively minor factor in
selection.
From the Department of Agronomy, University of Flor-
ida, Gainesville (Pfahler), and the North Florida Re-
search and Education Center, University of Florida,
Quincy (Barnett). This is Journal Paper No. 9407 of
the Florida Agricultural Experiment Station. We thank
Dr. Calvin Newton for supplying the seed of the short
culm population. Address reprint requests to Dr. Pfah-
ler, Department of Agronomy, 304 Newell Hall (IFAS
0311), University of Florida, Gainesville, FL 32611.
References
1. Hartl DL. Principles of population genetics. Sunder-
land, Massachusetts: Sinauer Associates; 1980. 488 p.
2. Mather K, and Jinks JL. Introduction to biometrical
genetics. Ithaca, New York: Cornell University Press;
1977. 231 p.
3. Schlegel R, Melz G, and Mettin D. Rye cytology,
cytogenetics and genetics—current status. Theor Appl
Genet 1986; 72:721-734.
4. Snedecor GW. Statistical methods, 5th ed. Ames,
Iowa: Iowa State University Press; 1956. 534 p.
Evidence for Apomixis in
Arabis
B. A. Roy and L. H. Rieseberg
The breeding system of a population of the
mustard Arabis holboellii var. retrofracta is
shown to be apomictic by means of enzyme
electrophoresis. Progeny tests revealed no
genetic variation within families, yet clonal di-
versity was relatively high. As there was no
evidence for recombination, it is suggested
that the populational variation was a result of
the polyphyletic origin of the apomictic clones.
Clonal organisms frequently bypass sex-
ual reproduction by means of modular
vegetative growth or by producing eggs
without undergoing meiosis (apomixis or
parthenogenesis). Evolutionary biologists
initially thought that organisms restricted
to asexual reproduction have limited evo-
lutionary potential.2515-26 More recently,
studies using molecular techniques have
shown that most populations of clonal or-
ganisms are more variable than was ex-
pected.8-16 The focus on genetic variation
in clonal plants has shifted to determining
how the variation is maintained.9122122
A few studies have examined genetic di-
versity in strictly apomictic plants using
evidence derived from enzyme electro-
phoresis.1012-14-23 Electrophoretic studies
allow the rapid identification and quanti-
fication of variation within and among
populations. Furthermore, by analysis of
progeny arrays, the frequency of recom-
bination or mutation can be estimated.8
Nevertheless, progeny testing has rarely
been employed in studies of apomictic
taxa.914
We analyzed isozyme variation in the
Downloaded from http://jhered.oxfordjournals.org/ at Universidad Nacional Autónoma de México on January 20, 2014
putative apomict Arabis holboelliiHornem.
var. retrofracta (Graham) Rydberg. Some
members of the A. holboellii complex have
characteristics suggestive of apomixis, in-
cluding the formation of pollen and em-
bryos with an unreduced chromosome
number.4 We specifically addressed three
questions: 1) Is this population of A. hol-
boellii var. retrofracta apomictic? 2) How
much isozyme diversity is there in a single
population? 3) At what rate are new iso-
zyme variants produced?
Materials and Methods
A 35-m transect was laid through a pop-
ulation of A. holboellii var. retrofracta in
Gunnison County, Colorado. At each even
meter mark, all the siliques present on the
nearest plant to the line were collected.
The seed from each plant was kept in in-
dividual packets, and each accession was
mixed thoroughly before planting so that
the seed represented a random choice. It
was not possible to analyze the parents
because A. holboellii var. retrofracta is often
monocarpic and because the parents were
dead at the time of seed collection. To
assess the amount of genetic diversity in
the population, we used one offspring from
each of the 30 individuals initially col-
lected for the population survey. The in-
dividual surveyed in the population anal-
ysis also was included in the progeny
arrays. We performed progeny analysis for
20 of the 30 parents. One family had only
16 individuals; otherwise, 24 siblings on
average were analyzed for each family.
We assayed the enzymes electrophoret-
ically on 12% starch gels. Phosphogluco-
isomerase (PGI), leucine aminopeptidase
(LAP), and triosephosphate isomerase
(TPI) were resolved on gel and electrode
buffer system 8 of Rieseberg and Soltis.19
System 1 of Soltis et al.24 was used to re-
solve isocitrate dehydrogenase (1DH) and
6-phosphogluconate dehydrogenase (6-
 common explanation for populational
variation in apomictic taxa is occasional
sexual reproduction.81124 However, our
4 progeny tests yielded no genetic variants,
i-i indicating that sexual reproduction oc-
curred rarely if at all in this population.
Three other mechanisms are frequently
hypothesized in the absence of sex: mu-
tation,810 migration from other popula-
tions where recombination does occur,13
and a polyphyletic origin of the polyploid
apomicts.3'11'25 It is not possible to rule out
migration at this time because it is not
II 4 known if sexual populations exist or if it
is likely for seed to have traveled from a

Downloaded from http://jhered.oxfordjournals.org/ at Universidad Nacional Autónoma de México on January 20, 2014
sexual population. However, a more prob-
able explanation can be inferred from our
data. The appearance of complete linkage
Figure 1. Photographs of starch gels of Arabis holboellii var. retrofracta stained for phosphoglucoisomerase
is expected for the progeny of an apomic-
(PGI). (A) Population survey of 30 individuals. Representatives of the four genotypes are shown at 1, 2, 3, and tic plant because of the lack of chromo-
4. (B) Array of 24 siblings of genotype 4, showing no variation within the family. somal segregation. Nevertheless, genetic
variants within apomictic populations
generally should be characterized by a
more random pattern of allelic associa-
tions if that variation has been produced
PGD). All enzyme stains followed Soltis et four phenotypes; thus, the percentage of by recombination. The presence of com-
al.,24 except LAP, which was modified by variation is 13.3%. pletely nonrandom allelic combinations
the addition of 30 mg of L-leucine-/3-naph- and the fact that only a small number of
thylamide. clones were detected suggest a polyphy-
Discussion
letic origin for these triploid clones.
The population survey showed variation
Results In conclusion, this population of A. hol-
among individuals (Figure 1A), yet all sib-
This population of A. holboellii var. retro- lings were invariant (Figure 1B). The most boelliivar. retrofracta is probably obligate-
fracta appears to be apomictic; there is likely explanation for this pattern is apo- ly apomictic. Genetic variation is present
variation in the population (Figure 1A) but mixis, as all progeny from an apomictic in the population but not within families.
not within families (Figure IB). The pop- individual should be genetically identi- The most likely explanation for the pop-
ulation survey of 30 individuals yielded cal.3'11 If recombination were occurring, ulational variation is polyphyletic clone
four genotypes when the banding patterns there would be segregation among prog- origin, which is indicated by the presence
for all five enzymes were combined. The eny, which clearly was not the case (Figure of nonrandom allelic associations and by
four genotypes each had a unique phe- IB). A less parsimonious alternative is the apparent absence of recombination.
notype for PGI (Figure 1A), TPI, and LAP. complete selfing, in which homozygous
Three phenotypes were observed for IDH, parents are expected to produce mono- From Rancho Santa Ana Botanic Garden, Claremont,
California (Roy and Rieseberg) and the Rocky Moun-
and two for 6-PGD. A highly nonrandom morphic offspring.1 Selfing is an unlikely tain Biological Laboratory, Crested Butte, Colorado
association of electromorphs was ob- explanation for the uniformity seen among (Roy). This study was supported in part by a Harriet
served (linkage disequilibrium); that is, Barclay Scholarship from the Rocky Mountain Biolog-
these progeny, because they all contained ical Laboratory to B.A.R. and NSF Grant BSR-87-22643
pattern 1 of PGI always occurred with pat- multibanded patterns. Such "fixed hetero- to L.H.R. We thank Paulette Bierzychudek for excellent
tern 1 of TPI, pattern 1 of LAP, and so forth. zygosity" could not arise from the selfing advice. Address reprint requests to B.A. Roy, Rancho
Santa Ana Botanic Garden, 1500 N. College Ave., Clare-
The number of loci and the inheritance of of homozygous parents unless gene du- mont, CA 91711.
the enzyme bands could not be deter- plications were present for every enzyme
mined, because testcrosses cannot be sampled. Even if this triploid was derived
made in apomicts and no closely related from highly divergent parents, a frequency
sexual taxa were available to aid interpre- of gene duplication this high is quite un-
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