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BEN # 223



BBBBB    EEEEEE   NN   N             ISSN 1188-603X
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BBBBB    EEEEE    NN N N             BOTANICAL
BB   B   EE       NN  NN             ELECTRONIC
BBBBB    EEEEEE   NN   N             NEWS

No. 223                              May 4, 1999

aceska@freenet.victoria.bc.ca        Victoria, B.C.
-----------------------------------------------------------
 Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2
-----------------------------------------------------------

HAEMOGLOBINS IN PLANTS
From: P.A. Guy <guy@cc.umanitoba.ca> and
      R.D. Hill <rhill@ms.umanitoba.ca>

Haemoglobin  is  most  commonly  known  as  a protein that binds
oxygen in the blood of animals and transports and  releases  the
oxygen  to  respiring tissues. However, haemoglobins are in fact
ubiquitous in most organisms, including not  only  animals,  but
also  bacteria,  protozoans, fungi and plants, where often their
functions have not been clarified.

Only in recent years has it  become  apparent  that  haemoglobin
proteins  probably  exist  in  all  plants. These can be broadly
grouped into symbiotic and nonsymbiotic haemoglobins. The  first
group  has  been  characterised  since the 1930s and include the
well studied `leghaemoglobins' found  in  the  root  nodules  of
nitrogen-fixing plants. Nonsymbiotic haemoglobins, however, were
only  identified  during  the late 1980s in the tree family, Ul-
maceae. They have since been studied in a range  of  dicots  in-
cluding  the  legumes;  monocot species from barley and wheat to
maize and rice and in the chloroplasts of algae.

All haemoglobins found across the spectrum of  living  organisms
can  reversibly  bind  oxygen.  Symbiotic  haemoglobins are syn-
thesised in proximity to the plant's symbiotic partners  and  in
large amounts. The protein readily gives up oxygen and acts very
well in transporting and regulating oxygen supply to the plant's
microorganism   associates.   Nonsymbiotic  haemoglobins  differ
markedly in gene homology  from  the  symbiotic  and  have  been
detected  in  a range of tissues, often only under stress situa-
tions and in much lower amounts than found for symbiotic haemog-
lobin. Also, they generally show remarkably  high  affinity  for
oxygen  in  that they will bind the molecule and keep it even at
very low cellular levels of oxygen. These  characteristics  dis-
count  a  role  in  facilitative oxygen transport and supply but
leaves the possibility of haemoglobin acting as an oxygenase  in
a  biochemical  reaction. The question which remains is "What is
their exact function?"

To this end the work on the  barley  haemoglobin  provides  some
insight.  It  is synthesised in flooded roots and can be induced
in isolated aleurone layers when atmospheric oxygen  is  reduced
to 5%. It is also induced in the normally germinating caryopsis,
in  both  the embryo and aleurone, and in the root and shoots of
the young seedlings. In aleurones it has been  shown  that  this
induction  is a consequence of lowered energy (i.e. ATP) levels.
It appears therefore that at least  barley  haemoglobin  is  re-
quired  during  times when energy demands of the cell can not be
met by the availability of oxygen for normal  respiration.  This
hypothesis  is  further  evident with the observation that maize
cells, genetically engineered to overproduce barley haemoglobin,
better maintain their energy levels when oxygen availability  is
compromised.  If  the  cells are engineered to eliminate haemog-
lobin, then they are unable to maintain energy levels under  the
same  conditions. This data provides valuable clues to the func-
tion and mode of action of nonsymbiotic haemoglobins in general.

The ubiquitous occurrence of nonsymbiotic haemoglobins in plants
indicates some fundamental roles in  plant  survival.  From  the
data for barley haemoglobin a role during transient waterlogging
or ice encasement can be envisaged, where oxygen availability is
low and there is a need to maintain energy levels. The plant can
resort  to  fermentation,  or  anaerobic  ATP production, but in
doing so produces toxic compounds such as  ethanol.  Haemoglobin
may  provide  an  alternative or complimentary route. This would
provide a plant able to synthesise haemoglobin with  a  distinct
selective  advantage.  As more data accumulates on other nonsym-
biotic haemoglobins, other roles may become apparent.

For more comprehensive reviews see:

Appleby, C.A. 1992. The origin and functions of haemoglobins  in
   plants. Sci. Prog. 76: 365-398.
Hill,  R.D.  1998. What are hemoglobins doing in plants? Can. J.
   Bot. 76: 707-712.


RE: BEN # 218 - SHOULD WE REJECT THE NAME 'SCIRPUS AMERICANUS'?
From: Adolf Ceska <aceska@victoria.tc.ca>

I was wrong to cite the ICBN [International  Code  of  Botanical
Nomenclature],  Article  57  of the Tokyo Code from 1993, when I
suggested that the name "Scirpus americanus" should be rejected.
At the time when  Schueler  made  the  typification  of  Scirpus
americanus, the ICBN did not have this article. The Seattle Code
(1969), effective at the time said: "A name is to be rejected if
it  is  used  in different senses and so has become a long- per-
sistent source of error." The Example  included  in  Article  69
implied  that you needed the name to be "applied almost equally"
to two different species before it was to be rejected.

The next version of the ICBN, the  Leningrad  Code  (1975),  was
already  phrased  differently. Again, Leningrad Code Article 69:
"A name must be rejected if it has been widely and  persistently
used  for  a  taxon  not including its type. Names thus rejected
shall be placed on a list of nomina rejicienda." Here  the  list
of  Nomina  Rejicienda  for species names was first established.
Had the Intermountain Flora (published in 1977 and probably  the
first  major  Flora that accepted the application of the name of
Scirpus americanus for "S. olneyi") followed that rule, it would
have used Scirpus pungens and Scirpus olneyi instead.

Retaining the name "Scirpus americanus"  will  lead  to  further
confusion, and the only logical way to minimize the damage is to
propose  that name for rejection. This action, however, requires
a formal proposal to the Nomenclatural Committee of the Interna-
tional Association of Plant Taxonomists. If we  reject  "Scirpus
americanus"  we  will  use  S. olneyi and S. pungens without any
confusion. At the same time, all  other  combinations  based  on
"Scirpus  americanus"  will  be rejected too. When we treat this
group of Scirpus s.lato as Schoenoplectus, we will use  Schoeno-
plectus olneyi (A. Gray) Palla and Schoenoplectus pungens (Vahl)
Palla.


IMPORTANT LITERATURE DATABASES, AGRICOLA AND AGRIS, NOW ONLINE
From: Scott Miller <smiller@icipe.org> originally posted
        on Entomo-l <entomo-l@listserv.uoguelph.ca>

Two major international databases of biological and agricultural
literature are now available for free on WWW:

AGRICOLA,  including both the journal article database from 1979
   to present, and the catalog of the USDA National Agricultural
   Library:
   http://www.nal.usda.gov/ag98/ag98.html

AGRIS, the FAO agricultural journal article database:
   http://bwg.fao.org/agrisnew/

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