Hugo K Dooner1, L.
Curtis Hannah2, and Shailesh Lal3
1 Waksman Institute, Rutgers University, Piscataway, NJ
08855
2 Horticultural Sciences, University of Florida,
Gainesville, FL 32611
3 Dept. of Biological Sciences, Oakland University,
Rochester, MI 48039
PREAMBLE: Helitrons are a novel class of
transposable elements discovered recently by computational analysis of the
complete genome sequences of C. elegans, Arabidopsis, and rice (Kapitonov, V. V. & Jurka, J. 2001.
Proc Natl Acad Sci USA
98, 8714-8719). It has become apparent that, in maize,
helitrons are both abundant and highly variable in sequence. There are currently no guidelines for
naming these elements, yet they are highly diverse in size and sequence because
they can pick up different gene fragments from the maize genome. Their diversity in sequence is
presently matched by their diversity in names. For example, they have been named according to the gene
where they insert (helitron sh2-7527; helitron ba1-ref) or the names of the locus and ferried gene fragments
(helitrons 9002NOPQ and 9008 HI) or identified with a letter (HelA; HelB).
In an attempt to introduce some order into this chaos, we would like to
suggest a nomenclature system for maize helitrons at the onset of the maize
genome sequencing project. These
guidelines arose from informal discussions at the Maize Genetics Conference in
Asilomar in March, 2006. We will
adopt these guidelines in our future publications and hope that other researchers
working with helitrons will adopt them, as well.
NOMENCLATURE:
Although they differ greatly in internal sequences, maize helitrons share
substantial sequence homology at their 5Õ and 3Õ ends, the latter being more
highly conserved. By comparing the
3Õ terminal 30 nucleotides of maize helitron sequences currently in the
database, it is clear that the elements group into two major clades, Hel1 and Hel2.
The elements in the larger Hel1 clade share at least 70% sequence identity; those in the
smaller Hel2 clade
are less related, sharing around 50% identity. We anticipate that many new elements will be identified as
helitrons because of conserved sequence features at their 5Õ and 3Õ ends (Kapitonov,
V. V. & Jurka, J. 2001). We
propose that new elements be grouped into either Hel1 or Hel2 superfamilies based on the relationship
of their 3Õ terminal 30 bp to the respective consensus sequences. Presently, the consensus sequences for Hel1 and Hel2 correspond to the sequences of HelA and HelB, respectively, in the bz locus of line McC (Lai, J., Li, Y.,
Messing, J. & Dooner, H. K. 2005. Proc Natl Acad Sci U S A 102, 9068-9073.). We also propose a criterion of 50% identity as the cutoff to
assign helitrons to a particular superfamily. If, by this criterion, additional helitron superfamilies are
identified in the future, they should be named Hel3, Hel4, and so on.
Following the
symbol designating the superfamily to which a helitron belongs would be a
number provided by a clearing house for helitron nomenclature (see below) and
an identifier, in parentheses, consisting of the locus or mutation where the
element is found, if known, separated by a colon from the name of the maize
line. For simplicity, the entire
helitron symbol should be italicized.
Thus, the helitron in sh2-7527 would be Hel1-1(sh2-7527), the first one discovered, and the one
in the bz genomic
region of McC would be Hel1-3(bz:McC). The NOPQ
element in locus 9002 of B73 would be Hel1-x(9002:B73), where x would stand for a number
assigned by the helitron nomenclature clearing house.
If a helitron is
discovered that is virtually identical to a previously described helitron, but
at a different locus, than the letter ÒaÓ is placed after the number assigned
to the first helitron and a letter ÒbÓ is placed after the number of the second
helitron. Again, the parenthetical
identifier would include the locus and line carrying the new helitron. For example, B73 has an almost
identical copy of Hel1-3(bz:McC) in chromosome 5S, at the same map location as umc1260.
The Hel1
element in the bz
locus would become Hel1-3a(bz:McC) and the one in 5S, Hel1-3b(umc1260:B73).
If a helitron is discovered that is virtually identical to a previously
described helitron at the same locus, but in a different line, then it should
be given the same designation as the first one, specifying in the parenthetical
identifier the names of the locus and line where found. For example, McC and W22 have a copy of
Hel1-3 at the same
site in the bz
genomic region. The helitron in
W22 would then be named Hel1-3a(bz:W22). Note that
this designation does not imply absolute sequence identity of the two helitrons
(which is, actually, not the case here).
As with genes, it will be up to the individual investigator to assess
sequence relatedness from the sequence database records. Finally, if a helitron is identified in
a BAC sequence not yet associated with any locus, the number of the BAC in the
GenBank record can substitute temporarily for the locus name.
Dr. Shailesh Lal
at Oakland University, MI, has agreed to serve as clearing house for assigning
blocks of numbers to investigators, institutions, or multi-institutional
projects, such as the maize genome initiative, that have identified new
helitrons.
AUTONOMOUS
HELITRONS. An autonomous
helitron has not been discovered.
Following maize genetics convention, an autonomous helitron can only be
defined by a functional test.
Therefore, helitrons should not be designated as ÒautonomousÓ solely on
the basis of sequence homology.
If, based on its sequence content, e.g., an intact replicase and
helicase, a helitron is considered to be potentially autonomous, it could be
called a putative autonomous helitron, yet given a symbol based on the general
nomenclature guidelines. It is
suggested that, once confirmed, an autonomous helitron be designated aHel, followed by an identifier as described
above.