The Tapetum and Systematics in Monocotyledons
          Carol A. Furness & Paula J. Rudall........................201


     Orbicules in Angiosperms Morphology, Function,
       Distribution, and Relation with Tapetum Types
          Suzy Huysmans, Gamal El-Ghazaly & Erik Smets..............240


     Robertsonian Fusion and Centric Fission in Karyotype
       Evolution of Higher Plants.
          Keith Jones...............................................273


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The Tapetum and Systematics In Monocotyledons

Carol A. Furness and Paula J. Rudall
Royal Botanic Gardens, Kew
Richmond, Surrey, TW9 3AE
UNITED KINGDOM
I. Abstract

This papercritically reviews the homologies and distribution of tapetum types
in monocotyledons, in relation to their systematics. Two main types of tapetum
are widely recognised: secretory and plasmodial, although intermediate types
occur, such as the 'invasive' tapetum described in Canna. In secretory tapeta,
a layer of cells remains intact around the anther locule, whereas in the
plasmodial type a multinucleate tapetal plasmodium is formed in the anther
locule by fusion of tapetal protoplasts. In 'invasive' tapeta, the cell walls
break down and tapetal protoplasts invade the locule without fusing to form a
plasmodium. When examining tapetum type it is often necessary to dissect
several developmental stages of the anthers. Secretory and plasmodial tapeta
are both widely distributed in monocotyledons and have probably evolved several
times, although there may be some systematic significance within certain groups.
Among early branching taxa, Acorus and Tofieldia have secretory tapeta, whereas
Araceae and Alismatales are uniformly plasmodial. The tapetum is most diverse
within Commelinanae, with both secretory and plasmodial types, and some
Zingiberales have an 'invasive' tapetum.  Lilianae (Dioscoreales, Liliales and
Asparagales) are almost uniformly secretory.

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Orbicules in Angiosperms Morphology, Function, Distribution, and Relation
    with Tapetum Types

S. Huysmans1 and E. Smets. Laboratory of Plant Systematics,
     Botanical Institute, K.U.Leuven, Kard. Mercierlaan 92,
     B-3001 Heverlee, BELGIUM

G. El-Ghazaly. Palynological Laboratory, Museum for Natural History,
     Roslagsvägen 101, S-10405 Stockholm, SWEDEN

1 Author to whom correspondence should be addressed.

I. Abstract
Orbicules or Ubisch bodies are sporopollenin particles lining the inne
tangential and sometimes also the radial tapetal cell walls. They only
occur in species with a secretory tapetum. The surface ornamentation of
orbicules and pollen of the same species is often strikingly similar.
Although orbicules are discovered more than a century ago, these structures
remain enigmatic since their function is still obscure. Proposed hypotheses
about their possible function are discussed. We also dealed with topics
such as the possible allergenicity of orbicules and their representation
in the fossil record. The use of orbicule characters for systematics is
reviewed.  The distribution of orbicules throughout the angiosperms, based
on a literature review from the first report until today, is shown in a list
with 314 species from 72 families. Those species found in the literature
without orbicules are presented together with their tapetum type. We
plotted this information on a dahlgrenogram to visualize the distribution
of orbicules. Orbicules occur in all subclasses of the angiosperms. Their
occurrence is not correlated with certain modes of pollination or habitats.

Résumé
Les orbicules ou corps d'Ubisch sont des particules de sporopollénine
couvrant la surface intérieure tangentiale et parfois la surface radiale
des cellules du tapétum. On ne les retrouve que dans les espèces possédant
un tapétum sécréteur. L'ornementation superficielle des orbicules et celle
du pollen d'une même espèce est souvent remarquablement similaire. Malgré
le fait que les orbicules ont été découvert il y a plus d'un siècle, ces
structures restent énigmatiques et leur fonction est toujours méconnue.
Les hypothèses proposées concernant la fonction éventuelle des orbicules
sont commentées dans cet article. Nous avons également traité des sujets
tels que les éventuels effets allergènes des orbicules ainsi que leur
présence dans les strates fossiles. L'utilisation de caractères orbiculaires
dans la systématique est analysée.  Nous présentons une liste de 314 espèces
appartenant à 72 familles possédant des orbicules, sur base d'une analyse
de la litérature à partir de la première observation jusqu'au présent. Pour
les espèces rapportées dans la litérature qui ne possèdent pas d'orbicules,
nous présentons aussi leur type de tapétum.  Nous avons projeté cette
information sur un Dahlgrenogramme afin de visualiser la distribution des
orbicules. Nous les retrouvons dans toutes les sous-classes des angiospermes.
Leur présence n'est pas correlée avec certains modes de pollinisation ou
avec divers types d'habitat.




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Robertsonian Fusion and Centric Fission in 
Karyotype Evolution of Higher Plants.


Keith Jones

 Jodrell Laboratory
Royal Botanic Gardens, Kew
Richmond, Surrey TW9 3DS
UNITED KINGDOM


I. Abstract

Robertsonian fusion and centric fission are uniquely detectable in comparative

studies of karyotype patterns. They are the most important types of karyotype

change in animals but seem to be relatively uncommon in higher plants.  Both

modify intra- and inter-chromosomal recombination and linkage relationships and

consequently patterns of genetic variation.  When differentiating populations or

species they can produce post-mating barriers to gene flow.  The number of

reported cases of fusion or fission in higher plants has increased over the years

but remains low, and most of these are casual comparisons of karyotypes without

any follow-up investigation.  This review focuses on more adequate studies made

in a few groups.
              

Studies in the Tradescantieae produce the strongest evidence for fusion as a 

type of ortho-selection in the subfamily.  Some species of Lycoris are also

considered to have evolved their karyotypes in that way.  Some genera of slipper

orchids and the cycad genus Zamia have populations where atypical chromosome

number increase can be attributed to fission probably as a result of stressful 

influences.

It is suggested that fusion may have been involved in the evolution of many

stable karyotypes and that fission is generally a secondary destabilizing

mechanism which may lead on to re-fusion in the long term.  Their proven

 incidence remains making it unwise to suggest that they have been major

influences in karyotype evolution in higher plants.

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Allen, A. 1977. Steps toward better scientific illustrations. Ed. 2. Allen Press, Lawrence, 
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Alston, R. E. 1968. The genetics of phenolic compounds. Pages 171-204 in J. B. Harborne (ed.), 
   Biochemistry of phenolic compounds.  Academic Press, New York.
CBE Style Manual Committee. 1983.  CBE style manual: A guide for authors, editors,  and 
   publishers in the biological sciences.  Ed. 5. Council of Biology Editors,  Bethesda, 
   Maryland.
Dahlgren, R. M. T., H. T. Clifford & F. F. Yeo. 1985.  The families of monocotyledons:  
   structure, evolution, and taxonomy.  Springer-Verlag, New York.
Funk, V. A. 1982.  Systematics of Montanoa Cerv. (Compositae).  Mem.  New York Bot. Gard. 
    36:1-133.
_____& D. R. Brooks. 198 1. Advances in cladistics.  The New York Botanical Garden, Bronx, 
   New York.
Gifford, E. M. & A. S. Foster. 1988.  Morphology and evolution of vascular plants. Ed. 3. 
   W. H. Freeman, New York.
Takhtajan, A. 1980.  Outline of the classification of flowering plants (Magnoliophyta). Bot.  
   Rev. 46:225-359.

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