IE83457B1 - Plants with modified flowers - Google Patents
Plants with modified flowersInfo
- Publication number
- IE83457B1 IE83457B1 IE1990/2911A IE291190A IE83457B1 IE 83457 B1 IE83457 B1 IE 83457B1 IE 1990/2911 A IE1990/2911 A IE 1990/2911A IE 291190 A IE291190 A IE 291190A IE 83457 B1 IE83457 B1 IE 83457B1
- Authority
- IE
- Ireland
- Prior art keywords
- promoter
- plant
- dna
- cells
- gene
- Prior art date
Links
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Classifications
-
- C—CHEMISTRY; METALLURGY
- C07—ORGANIC CHEMISTRY
- C07K—PEPTIDES
- C07K14/00—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof
- C07K14/195—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from bacteria
- C07K14/32—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from bacteria from Bacillus (G)
-
- C—CHEMISTRY; METALLURGY
- C07—ORGANIC CHEMISTRY
- C07K—PEPTIDES
- C07K2319/00—Fusion polypeptide
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8287—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for fertility modification, e.g. apomixis
- C12N15/8289—Male sterility
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
- C12N9/0004—Oxidoreductases (1.)
- C12N9/0089—Oxidoreductases (1.) acting on superoxide as acceptor (1.15)
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
- C12N9/10—Transferases (2.)
- C12N9/1003—Transferases (2.) transferring one-carbon groups (2.1)
- C12N9/1007—Methyltransferases (general) (2.1.1.)
Description
PATENTS ACT, 1992
2911/90
PLANTS WITH MODIFIED FLOWERS
AVENTIS CROPSCIENCE N.V.
i83457
This invention relates to a method of restoring
fertility to a transgenic nuclear male—sterile or
female—sterile plant by crossing such a sterile plant
with a transgenic fertility—restorer plant to provide a
transgenic fertility—restored plant having’ a foreign
DNA sequence from the nuclear genome of the restorer
plant that is stably integrated into the nuclear genome
of the restored plant. The foreign DNA sequence of this
foreign DNA
that: 1)
invention contains a (hereinafter the
"fertility—restorer DNA") encodes a first
protein. or polypeptide which is an inhibitor of a
ribonuclease and which, when produced or overproduced
in a cell of a flower, particularly a male or female
reproductive organ thereof,
or a seed or an embryo of
restored plant, prevents the activity in the
flower, seed or embryo cell of a second. protein or
polypeptide which is the ribonuclease and which, when
produced or overproduced in the flower,
cell,
the metabolism,
seed or embryo
would otherwise significantly disturb adversely
functioning and/or development of the
and. 2) is in the same
flower, seed or embryo cell;
transcriptional unit as, and under the control of, a
first promoter which is capable of directing expression
of the fertility—restorer DNA at least in the same
flower or seed or embryo cells of the restored plant
where the second RNA, protein or polypeptide is being
The second,
"sterility—DN "
produced or overproduced. protein or
polypeptide is encoded by a foreign
that is from the nuclear genome of the sterile plant,
that is also stably integrated into the nuclear genome
of the restored plant and that is under the control of
capable of: i)
a "sterility—promoter“ which is
directing expression of the sterility DNA selectively
in specific cells of each flower, particularly at least
one ‘male or at least one female reproductive organ
thereof, or each seed or each embryo of the restored
plant and ii)
male- or female-sterile in the absence of expression of
thereby rendering the restored plant
the fertility-restorer DNA in the specific flower, seed
or embryo cells.
The foreign DNA of this
transferred from the restorer plant into the restored
sequence invention,
plant, is optionally a foreign chimaeric DNA sequence
that can also contain a second foreign DNA (the “first
marker DNA“) that:
polypeptide which, when present at least in a specific
of the plant,
entire plant easily separable or distinguishable from
) encodes a third RNA, protein or
tissue or specific cells renders the
other plants that do not contain the third RNA, protein
or polypeptide at least in the specific tissue or
cells of the plant: 2) is
and under the control of, a
specific in the same
transcriptional unit as,
second promoter which is capable of directing
expression of the first marker DNA in at least specific
tissue or specific cells of the plant; and 3) is in the
same genetic locus of the nuclear genome of the
restored plant as the fertility-restorer DNA.
This invention also relates to a foreign chimaeric
DNA sequence that contains at least one fertility-
restorer DNA under the control of at least one first
promoter and that can also contain, adjacent to the
fertility-restorer DNA(s) and the first promoter(s), at
least one first marker DNA under the control of at
least one second promoter.
This invention further relates to: a vector that
contains the foreign DNA sequence of this invention and
is suitable for the transformation of a plant cell,
whereby the foreign DNA sequence is stably integrated
the foreign DNA sequence,
into the nuclear genome of the cell: the resulting
fertility-restorer plant cell; cultures of such
fertility-restorer plant cells; a fertility-restorer
plant and its reproductive material (e.g., seeds) which
can be regenerated from such a fertility-restorer plant
stably
sequence 7 a
cell and the nuclear genome of which contains,
the foreign DNA
fertility-restored plant and its reproductive material
integrated therein,
containing, stably integrated in their nuclear genome,
together with at least one
sterility DNA under the control of at least
and a cell of the
restored plant, as well as cultures thereof.
sterility promoter; fertility-
This invention yet further relates to a process
for producing the restorer plant and its reproductive
material by transforming a cell of the plant with the_
foreign DNA sequence whereby the fertility-restorer DNA
is: 1) under the control of the first ‘promoter' and
optionally in the same genetic locus as the first
marker DNA under the control of the second promoter;
and 2) stably integrated into the nuclear genome of the
plant's cells.
The invention further relates to hybrid seeds
produced by crossing: 1) the restorer plant, preferably
also containing, its nuclear
the
conferring a resistance to a herbicide on the restorer
stably integrated in
genome, first marker DNA encoding a protein
plant; with 2) a nuclear male- or female-sterile plant
which has, stably integrated in its nuclear genome a)
the sterility DNA under the control of the sterility
promoter and, adjacent to the sterility DNA, preferably
within the same genetic locus of the nuclear genome, b)
a second marker DNA, encoding a fourth RNA, protein or
polypeptide and preferably also conferring a herbicide
resistance on the sterile plant, under the control of a
third promoter capable of directing expression of the
second ‘marker DNA in at least a specific tissue or
specific cells in which expression of the second marker
DNA renders the plant easily separable or
distinguishable from those in which there is not such
expression. This invention particularly relates to such
hybrid
preferably ix: a substantially random population, with
seeds as produced on a commercial scale,
increased efficiency of cross-pollination and without
the need for extensive hand-labor.
Background of the Invention
Hybridization of plants is recognized as an
important process for producing offspring having a
combination of the desirable traits of the parent
plants. The resulting hybrid offspring often have the
ability to outperform the parents in different traits,
yield, adaptability to
changes, and disease resistance. This ability is called
“hybrid
hybridization has been used extensively for improving
such as in environmental
"heterosis' or vigor". As a result,
major crops, such as corn, sugarbeet and sunflower. For
a number of reasons, primarily related to the fact that
most plants are capable of undergoing both self-
cross-pollination, the
cross-pollination of plants without significant self-
pollination and controlled
pollination, to produce a harvest of hybrid seeds, has
been difficult to achieve on a commercial scale.
In nature, the vast majority of crop plants
produce male and female reproductive organs on the same
plant, usually in close proximity to one another in the
same flower. This favors self-pollination. some plants,
however, are exceptions as a result of the particular
morphology of their reproductive organs which favors
cross-pollination. These hybrid
plants produce
offspring with improved vigor and adaptability. one
such morphology in Cannabis ggp; (hemp) involves male
and female reproductive organs on separate plants.
Another' such morphology in Zea mays (corn) involves
male and female reproductive organs on different parts
of the same plant. Another such morphology in Elaeis
guineensis (oilpalm) involves male and fertile female
gametes which become fertile at different times in the
plant's development.
Some other plant species, such as Ananas comosus
(pineapple), through the
particular physiology of their reproductive organs.
Such "self-
incompatibility system" whereby the pollen of one plant
favor cross-pollination
plants have developed a so—cal1ed
is not able to fertilize the female gamete of the same
plant or of another plant with the same genotype.
isome other plant species favor cross-pollination
so-called genomic
this
characteristic, the plants‘ anthers degenerate before
by naturally displaying the
characteristic of "male-sterility". By
pollen, produced by the anthers. reaches maturity. See:
"Male-Sterility in Higher Plants", K.L.H. Kaul, 1987,
in: Monographs on Theoretical and Applied Genetics 10,
Edit.
characteristic is believed to result from a wide range
Springer Verlag. Such a natural male-sterility
of natural mutations, most often. involving
deficiencies, and this characteristic can not easily be
maintained in plant species that predominantly self-
pollinate, since under natural conditions, no seeds
will be produced.
Some types of naturally occurring male-sterility
are cytoplasmatically encoded, while others are nuclear
encoded. One type of male-sterility is the result of a
combination of both nuclear encoded male-sterility and
The male-
cytoplasmatically encoded .male-sterility.
sterility inducing nuclear alleles are usually
recessive, and only plants that contain the male-
sterility cytoplasmic allele and that are homozygous
for the male-sterility inducing nuclear allele are
phenotypically male-sterile. In this type of plant,
corresponding dominant male-fertility inducing alleles
or "fertility male—fertile
restorers" produce a
phenotype. As a result, the male-sterile offspring of
this type of plant can be made male-fertile by
pollinating the male-sterile plants with pollen
containing the fertility restorers . As a result, the
offspring of plants of this type are of commercial
value where the economic product is seeds (e.g., for
plants such as corn, sorghum and sunflower).
Host of the
sterility genes and their
restorer genes have not been used
known naturally occurring male-
corresponding fertility-
in breeding or
essentially two
the
for the male-sterility and restoration
and b) low
capability of the crops in which they occur.
production of new varieties for
insufficient quality of genes
reasons: a)
responsible
characteristics; cross-pollination
. The quality of the genes
To realize the full potential of a male—sterility/
fertility—restorer system, several quality
requirements have to be achieved:
a) Stability of the genes encoding the male—sterility
broad
conditions.
under a range of different environmental
conditions. Most of the currently known
systems,whether they are nuclear or cytoplasmatically
encoded, do not display sufficient stability. As a
consequence of this, under some unpredictable
climatological conditions, self—pollination occurs
within the plants, and heterogeneous offspring are
harvested. According to seed certification
requirements, not more than 1% of non-hybrid seed
is tolerated for most major field crops.
No side effects on the plants. Many cytoplasmic
male—steri1ity genes induce a decrease in plant
This can be tolerated up to a certain
if the hybrid effect
significant improvement of the crop compared to
vigor.
level, vigor offers a
the negative effect. Another side effect which has
been observed in crops carrying male-sterility
genes consists of an enhanced sensitivity to some
(9I
cytoplasmic male-sterility are highly susceptible
plant pathogens corn plants carrying T-
to Helminthosporium maydis infections).
Restorer genes also often display negative side
effects although these are usually not due to the
genes themselves but to genes closely linked to
the restore: genes. These side effects consist, in
most cases, of an increased disease or pest
susceptibility or a decreased quality of the crop.
Efficiency of cross-pollination
Reasonably efficient cross-pollination is
essential for the production of hybrid seeds at an
acceptable cost. For major field crops that are
poorly adapted to
unrealistic to assure cross—pollination by hand.
cross-pollination, it is
it has been envisaged to sell, as a
not the F1 hybrid, but the
(e.g. , cotton and
The disadvantage of this method lies,
Therefore,
commercial product,
selfed F2
wheat).
however,
and the
combinations. To assure high yield of a crop by a
offspring thereof
in the loss of homogeneity and heterosis
segregation of specific useful gene
farmer, it is advantageous that hybrid crops be
fully fertile (with the very
efficient cross—pollinating species such as corn
exception of
and oilseed rape). This is particularly the case
with crops that form heavy or sticky pollen which
is not easily transported by wind (e.g., cotton),
with crops that are not attractive to pollinating
insects (e.g., wheat) and with crops which display
cleistogamy (e.g., soybean).
Detailed Description of the Invention
In accordance with this invention, a fertility-
restorer plant is produced from a single cell of a
plant by transforming the plant cell in a well known
manner to stably insert, into the nuclear genome of the
cell, the foreign DNA sequence of this invention. The
foreign DNA sequence comprises at least one fertility-
restorer DNA that is under the control of, and fused at
its 5' end to, the first promoter and is fused at its
3' end to suitable termination (or
regulation) signals,
signal. Thereby, the first RNA, protein or polypeptide
is produced or overproduced in cells of at least each
transcription
including a polyadenylation
of the restorer plant's flowers, preferably one or more
male or one or more female reproductive organs thereof,
and/or seeds and/or embryos, so that when the restorer
plant is crossed with a nuclear male-sterile or nuclear
female—sterile hybrid male-fertile female-
fertile obtained. The DNA
sequence can also comprise at least one first marker
plant,
offspring are foreign
DNA that is under the control of, and is fused at its
' end to, the second promoter and is fused at its 3'
suitable transcription termination signals,
The first marker
end to
including a polyadenylation signal.
DNA is preferably in the same genetic locus as the
fertility-restorer DNA, whereby the third RNA, protein
or polypeptide is produced in at least the specific
tissue or specific cells of the fertility-restorer
plant so that the plant can be easily distinguished
and/or separated from other plants that do not contain
the third RNA, protein or polypeptide in the specific
tissue or specific cells. This guarantees, with a high
degree of certainty, the joint segregation of both the
fertility—restorer DNA and the first marker DNA into
offspring of the plant.
The cell of a plant (particularly a plant capable
Agrobacterium)
transformed in accordance with this invention, using a
of being infected with is preferably
vector that is a disarmed Ti-plasmid containing the
foreign DNA sequence and carried by Agrobacterium. This
transformation can be carried out using procedures
described, for example, in European patent publications
,116,718 and 0,270,822. Preferred Ti-plasmid vectors
contain the foreign DNA sequence between the border
sequences, or at least located to the left of the right
of the T-DNA of the Ti-plasmid. Of
course, other types of vectors can be used to transform
the plant cell,
transfer
border sequence,
using procedures such as direct gene
described, for in European
0,223,247),
for example,
((as example,
patent publication pollen mediated
transformation (as described, in European
patent publication 0,270,356, PCT
W085/01856, and European patent publication 0,275,069),
(as described for
publication
in vitro protoplast transformation
4,684,611),
mediated transformation (as described,
example, in US patent plant RNA virus-
for example, in
and Us patent
and liposome-mediated transformation (as
European patent publication 0,067,553,
4,407,956)
described, for example, in US patent 4,536,475).
Preferably, a fertility-restorer plant of this
invention is provided by transforming a plant cell with
a disarmed Ti-plasmid vector containing the foreign DNA
sequence with a DNA under the
control of a first promoter and optionally a first
marker DNA under the control of a second promoter. The
marker DNA can be upstream or downstream of the
fertility-restorer DNA in the Ti-plasmid vector, but
preferably, the two are adjacent to one another and are
fertility—restorer
located between the border or at least
located to the left of the right border sequence of the
Ti-plasmid that they
transferred together into the nuclear’ genome of the
plant cell. However, if desired, the cell can initially
be transformed with the foreign DNA sequence containing
the fertility—restorer DNA and the first promoter and
can subsequently be transformed with the marker DNA and
sequences
vector, so are properly
the second promoter, inserted into the same genetic
locus in the cell's nuclear genome as the fertility-
restorer DNA, or this transformation can be carried out
vice versa. Suitable vectors for this purpose are the
same as those discussed above for transforming cells
with the foreign DNA sequence. The preferred vector is
a disarmed Ti-plasmid vector.
The selection of the fertility-restorer DNA of
this invention is not critical but is dependent on the
selection of, and must correspond to, the sterility DNA
which is responsible for the male- or female-sterility
characteristic to be restored. In particular, the
production or overproduction of the first RNA, protein
or' polypeptide encoded by the fertility-restorer DNA
has to neutralize, block, offset, overcome or otherwise
prevent the specific activity of the second protein or
polypeptide which is a ribonuclease encoded by the sterility
DNA in flower cells, preferably cells of at least one male or
at least one female reproductive organ, or in seed
of the
Examples of male- and female- sterility DNAs, to which
the fertility-restorer DNAs of this invention must
correspond, and the which they
counteract, are described in European
,344,029 and 0,412,006,
which are incorporated herein by reference. A suitable
fertility-restorer DNA can be selected and isolated in
cells or in embryo cells restored plant.
action of must
patent
publications respectively,
a well-known manner to overcome the effects of the
sterility DNA in any cell of a flower, particularly a
male or female organ, a seed and/or an embryo, in which
the sterility promoter causes the sterility DNA to be
expressed.
Preferred examples of fertility-restorer DNAs
encode: barstar which neutralizes the activity of
barnase (which degrades RNA molecules by hydrolyzing
the bond after any guanine residue).
Still further examples of fertility-restorer DNAs
can be combinations of one or more of the different
fertility-restorer DNAs cited above.
By "foreign" with regard to the foreign DNA
sequence of this invention is meant that the foreign
DNA sequence contains a foreign fertility—restorer DNA
and/or a foreign first promoter. By with
regard to a DNA, such as a fertility-restorer DNA and a
"foreign"
first promoter, as well as a first marker DNA, a second
promoter and any other DNA in the foreign DNA sequence,
is meant that such a DNA is not in the same genomic
environment in a plant cell, transformed with such a
DNA in accordance with this invention, as is such a DNA
when it is naturally found in the cell of the plant,
or the like,
This means,
bacteria, animal, fungus, virus, from
which such a DNA originates. for example,
that a foreign fertility-restorer DNA or first marker
DNA can be: 1) a nuclear DNA in a plant of origin; 2)
endogenous to the transformed plant cell (i.e., from a
plant of origin with the same genotype as the plant
within the
transcriptional unit as its own endogenous promoter and
being transformed); and 3) same
3' end transcription regulation signals (from the plant
of this
in the transformed plant cell; but 4)
of origin) in the foreign DNA sequence
invention
inserted in a different place in the nuclear genome of
the transformed plant cell than it was in the plant of
origin so that it is not surrounded in the transformed
plant cell by the genes which surrounded it naturally
in the plant of origin. A foreign fertility-restorer or
first marker DNA can also, for example, be: 1) a
nuclear DNA in a plant of origin; and 2) endogenous to
the transformed plant but 3) in the
transcriptional unit as a different (i.e., not its own)
cell: same
endogenous promoter and/or 3' end transcription
regulation signals in a foreign chimaeric DNA sequence
of this
foreign fertility-restorer or first marker DNA can
invention in a transformed plant cell. A
for example, be:
origin: and 2)
cell: but 3)
heterologous
also, 1) a nuclear DNA in a plant of
endogenous to the transformed plant
in the same transcriptional unit. as a
and/or 3'
regulation signals in a foreign chimaeric DNA sequence
of this
foreign fertility-restorer or first marker DNA can
promoter end transcription
invention in a transformed plant cell. A
also, for example, be heterologous to the transformed
plant cell and in the same transcriptional unit as an
endogenous promoter and/or 3' transcription regulation
signals (e.g., from the nuclear genome of a plant with
the same genotype as the plant being transformed) in a
foreign chimaeric DNA sequence of this invention in a
cell. An foreign
fertility-restorer DNA could come from the nuclear
transformed plant example of a
genome of a plant with the same genotype as the plant
transformed and encode an inhibitor of a
that
being
ribonuclease, is endogenous to the plant being
transformed, so that the enzyme is overproduced in
transformed cells in order to neutralize the activity
of a ribonuclease (i.e., a second protein encoded by a
male— or female—sterility DNA) which would
significantly disturb adversely the metabolism,
functioning and/or development of flower cells,
particularly male or female organ cells, or seed cells
or embryo cells, in which such an enzyme is expressed.
Preferably, each fertility—restorer DNA and first
marker DNA is heterologous to the plant cell being
transformed.
By "heterologous" with regard to a DNA, such as a
fertility-restorer DNA,
first marker DNA and any other DNA in the foreign DNA
sequence of this invention, is meant that such a DNA is
a first or third promoter, a
not naturally found in the nuclear genome of cells of a
plant with the
Examples
same genotype as the plant being
DNAs
chloroplast and mitochondrial DNAs obtained from a
plant with the
but preferred examples are chloroplast,
transformed. of heterologous include
same genotype as the plant being
transformed,
mitochondrial,
different genotype than the plant being transformed,
and nuclear DNAs from plants having a
DNAs from animal and bacterial genomes, and chromosomal
and plasmidial DNAs from fungal and viral genomes.
By "chimaeric" with regard to the foreign DNA
sequence of this invention is meant that at least one
of its fertility-restorer DNAs: 1) is not naturally
found under the control of its first promoter for the
one fertility-restorer DNA: and/or 2) is not naturally
found in the same genetic locus as at least one of its
first marker DNAs. Examples of foreign chimaeric DNA
sequences of this invention comprise: a fertility-
restorer DNA of bacterial origin under the control of a
first promoter of plant origin: and a fertility-
restorer DNA of plant origin under the control of a
first promoter of plant origin and in the same genetic
locus as a first marker DNA of bacterial origin.
By "flower" is meant to include the entire shoot
axis, sepals, petals, male reproductive organs (or
stamens) and/or female reproductive organs (or carpels)
whose wholly or partly, retarded or arrested
development would prevent the development and/or
propagation of viable seeds in the flower or the
development and/or propagation of its male gametes; by
"male organ" or "male reproductive organ‘ is meant the
entire organ of a flower that is involved in the
production of the male gamete, as well as one or more
of its individual parts such as its anther, pollen and
filament; and by "female organ" or "female reproductive
organ" is meant the entire organ of a flower that is
involved in the production of the female gamete and/or
viable seeds and/or viable embryos, as well as one or
more of its individual parts such as its ovary, ovule,
style, stigma, corolla, disc, septum, callyx and
placenta. By "embryo" is meant to include the entire
embryo of a plant, as well as one or ‘more of its
individual parts such as its embyo axis and embryo
cotyledons.
So that the fertility-restorer DNA is expressed in
at least those specific cells of a fertility-restored
plant in which the sterility DNA is expressed, it is
preferred that the
expression of the fertility-restorer DNA, be a promoter
first promoter, which controls
capable of directing gene expression in at least the
same fertility-restored plant cells (i.e., the specific
flower cells, preferably male or female organ cells, or
seed cells or embryo cells), in which the sterility DNA
is selectively expressed under the control of the
sterility promoter. Such a first promoter can be an
endogenous promoter or an exogenous promoter and can be
from the nuclear genome or from the ntochondrial or
chloroplast genome of a plant cell. In any event, the
first promoter is foreign to the nuclear genome of the
plant cell being transformed. The first promoter can be
a constitutive promoter but can also be the same
selective promoter as the sterility promoter.
Preferably, the first promoter causes the restoration
of. fertility through the
production of at least
sufficient amounts of fertility-restoring first RNA,
protein or polypeptide selectively in the same specific
flower, seed or embryo cells, particularly in the same
specific flower cells, as those in which the sterility
DNA is expressed.
The first promoter of this
selected and isolated 5J1 a known manner from a plant
species, for example as described in: European patent
Publication 0,344,029 which is incorporated herein by
reference and which discloses a male-sterility promoter
that directs expression of a sterility DNA selectively
in stamen (e.g., cells of a plant and is
effective to prevent expression of the sterility DNA in
other parts of the European patent
publication 0,412,006
herein by reference and which discloses a female-
sterility promoter that directs expression of a
sterility DNA selectively in
particularly cells of a female organ (e.g., pistil), or
seed cells or embryo cells of a plant and is effective
to prevent expression of the sterility DNA in other
parts of the plant. For example, a suitable endogenous
first promoter can be
invention can be
anther)
plant; and
which is also incorporated
cells of flowers,
organ- or tissue—specific
identified and isolated in a plant, by:
. searching for an mRNA which is only present in the
plant during the development of its flowers, seeds
or embryos, preferably its anthers, pollen,
filaments, ovary, ovule, style, stigma, placenta,
calyx, scutellum, septum, seedcoat, endosperm or
embryo cotyledons: V
. isolating this specific mRNA:
. preparing a cDNA from this specific mRNA:
.4. using this cDNA as a probe to identify the regions
in the plant genome which contain DNA coding for
the specific mRNA: and then
. identifying the portion of the plant genome that
is upstream (i.e., 5') from the DNA coding for the
specific mRNA and that contains the promoter of
this DNA.
The genes controlled by these first promoters can
further be used as probes as in step 4, above. Under
hybridizing conditions, such a probe will hybridize to
DNA coding for a specific mRNA in a mixture of DNA
sequences from the genome of another plant species
(Maniatis et al (1982) Molecular Cloning. A Laboratory
Manual. Ed. Cold Spring Harbor Laboratory). Thereafter,
as in step 5 above, a specific first promoter for
another plant species can be identified.
Examples of’ male organ-specific first promoters
the PTA29 promoter, the
PTA26 promoter and the PTA13 promoter, as described in
European patent publication 0,344,029, which have been
and sterility promoters are:
isolated from tobacco and are tapetum—specific
promoters; as well as any promoter of a gene encoding a
tapetum—specific mRNA hybridizable to the genes TA29,
TA26 or TA13 of European patent publication 0,344,029,
from which genes the PTA29, PTA26 and PTA13 promoters
have been isolated. Examples of female organ-specific
first promoters and sterility promoters are: the style
and/or PSTMGO7,
PSTMG08, PSTMG4B12 and PSTMGBC9, and the ovule—specific
promoter corresponding to the cDNA clone pMON9608 as
stigma—specific promoters, such as
described in European patent publication 0,412,006; as
well as a promoter of a gene encoding i) a style—stigma
specific or ii) an ovule—specific mRNA. hybridizable
respectively to jj a STMG—type style—stigma specific
gene or ii) CDNA. clone pMON9608 of European. patent
publication 0,412,006.
If more than one nuclear sterility DNA is present
in the transgenic sterile plant which is to be crossed
with the transgenic fertility-restorer plant of this
invention, the restorer plant may need to have inserted
into its nuclear genome more than one fertility-
restorer DNA of this invention, corresponding in number
at least to the number of sterility DNAs in the nuclear
genome of the sterile plant. All the fertility-restorer
DNAs can be under the control of a single first
each fertility-restorer DNA
promoter, but preferably,
is under the control of its own separate first
promoter, which. will direct expression of the first
RNA,
where the sterility promoters cause the sterility DNAs
protein or polypeptide at least in those cells
to express the second RNA, protein or polypeptide. Each
fertility—restorer DNA is
and all the fertility-restorer DNA(s)
their first promoter(s) are preferably adjacent to one
adjacent to its first
promoter, and
another in the foreign DNA sequences of this invention
and in any vectors used to transform plant cells with
foreign DNA
necessary that the fertility-restorer DNAs be adjacent
such sequences. However, it is not
to one another in the foreign DNA sequence, and in some
cases, they may be inserted into the nuclear genome of
the restorer plant through independent transformation
events.
The selection of the first marker DNA of this
invention also is not critical. A suitable first marker
DNA can be selected and
so that it encodes the third RNA,
expressing the first
isolated in a well known
manner, protein or
polypeptide that allows plants,
marker DNA, to be easily distinguished and separated
from plants not expressing the first marker DNA. In
the first marker DNA encodes the same RNA,
protein or polypeptide as the second marker DNA encodes
many cases,
in the nuclear male- or fema1e—sterile plant, the
fertility of which is to be restored in accordance-with
this invention. Examples of the first marker DNAs are
the marker DNAs in the nuclear genomes of the nuclear
male— and female—sterile plants described in European
patent publications 0,344,029 and 0,412,006 which
encode proteins or polypeptides that: provide a
distinguishable color to plant cells, such as the Al
gene encoding dihydroquercetin—4—reductase
(1987) Nature 330, 677-678)
(1988)
);
(Meyer et al
and the glucuronidase gene
(Jefferson. et al Proc. Natl. Acad. Sci. USA
(”PNAS”) 83, provide a specific morphological
characteristic to a plant such as dwarf growth or a
different shape of the leaves; confer on a plant stress
tolerance, such as is provided by the gene encoding
superoxide dismutase as described in European patent
88402222.9 which is the
application priority
application claimed in
,359,617,
European patent publication
confer disease or pest resistance on a
plant, such as is provided by a gene encoding a
Bacillus thuringiensis endotoxin conferring insect
resistance as described in European patent publication
0,193,259; or confer on a plant a bacterial resistance
such as is provided by the bacterial peptide described
in European patent publication 0,299,828.
Preferred first marker DNAS encode third proteins
or polypeptides inhibiting or neutralizing the activity
of herbicides such as: the sfr gene and the sfrv gene
encoding enzymes conferring resistance to glutamine
synthetase inhibitors such as Bialaphos and
phosphinotricine as described in European patent
publication 0,242,246; and genes encoding modified
target enzymes for certain herbicides that have a lower
affinity‘ for the herbicides than naturally" produced
endogenous enzymes, such as a modified glutamine
synthetase as target for phosphinotricine as described
in European patent publication 0,240,792 and a modified
—enolpyruvylshikimate—3 phosphate synthase as a target
for glyphosate as described in European patent
publication 0,218,571. other first marker DNAs encode
third proteins which neutralize the action of the
herbicide bromoxynil (Stalker et al (1988) in: Genetic
Improvements of Agriculturally important Crops. Ed:
R.T. Fraley, N.M. Schell.
Harbor Laboratories) or the herbicide sulfonylurea (Lee
et al (1988) EMBO J. 1, 1241-1248) or the herbicide 2,4
D (disclosed at
Plant Molecular
1988).
The second
Frey and J. Cold Spring
the 2nd International Symposium of
Biology, Jerusalem, 13-18 November
of this which
controls the first marker DNA, can also be selected and
promoter invention,
isolated in a well known manner so that the first
marker DNA is expressed either selectively in one or
tissues or specific cells or
more specific
constitutively in the entire plant, as desired
depending on the nature of the third RNA, protein or
polypeptide. In many cases, the second promoter is the
same as the third promoter which controls the second
marker DNA in the male- or female-sterile plant, the
fertility of which is to be restored in accordance with
if the first marker DNA
it may be useful to
this invention. For example,
encodes an herbicide resistance,
have the first marker DNA expressed in all cells of the
plant, using a strong constitutive second promoter such
(Odell et al (1985) Nature ‘Q13,
(Hull and Howell (1987)
the promoter of the nopaline
as a 35S promoter
810-812),
Virology gg,
a 35S'3 promoter
432-493),
synthetase gene ("PNOS”)
Estrella (1983) Nature ggg, 209-213) or the promoter of
the octopine synthase gene ("POCS" [De Greve et al
(1982) J. Mol. Appl. Genet. 1 (6), 499-511]). If the
first marker DNA encodes a protein conferring disease
of the Ti-plasmid (Herrera-
resistance, it may be useful to have the first marker
DNA selectively expressed in wound tissue by using, for
example, a second promoter which is a TR promoter such
as the TR1' or TR2' promoter of the Ti-plasmid (Velten
et al (1984) EMBO J. ;, 2723-2730). If the first marker
DNA encodes a herbicide resistance, it may be useful to
have the first marker DNA selectively expressed in
green tissue by using as the second promoter, for
example, the promoter of the gene encoding the small
subunit of Rubisco (European publication
0,242,246).
it may be useful to select the second promoter so that
patent
_If the first marker DNA encodes a pigment,
the first marker DNA is expressed in specific cells,
leaf
preferably in the outer layer of the seed coat.
such as petal cells, cells or seed cells,
one can identify and isolate in a well known
manner a tissue-specific second promoter, suitable for
inclusion in the foreign DNA sequence of this invention
in a restorer plant or a restored plant of this
invention, whereby the plant can be easily
distinguished as carrying the first marker DNA under
the control of the second promoter. This can be done
by:
. searching for an mRNA which is only present in the
plant during the development of a specific tissue,
such as its petals, leaves or seeds:
. isolating this tissue-specific mRNA:
. preparing a cDNA from this tissue-specific mRNA:
. using this cDNA as a probe to identify the regions
in the plant genome which contain DNA coding for
the tissue-specific mRNA; and then
. identifying the portion of the plant genome that
is upstream from the DNA coding for the tissue-
specific mRNA and that contains the promoter for
said DNA.
If more than one first marker DNA is present in
all the
first marker DNAs can be under the control of a single
the foreign DNA sequence of this invention,
second promoter, but preferably, each first marker DNA
is under the control of its own separate second
promoter. More preferably, each first. marker DNA is
under the control of its own
encodes a different third RNA, protein or polypeptide,
second promoter and
providing different distinguishable characteristics to
a transformed plant. the first marker
DNA(s)
each other and to the one or more fertility-restorer
In any event,
and second promoter(s) should be adjacent to
DNAs contained in the foreign DNA sequence of this
invention and in any vector used to transform plant
cells with the foreign DNA sequence.
It is generally preferred that the first RNA,
encoded by the fertility-
restorer DNA, substantially prevents the activity of the
second RNA, protein or polypeptide,
sterility DNA,
plant cells in which the sterility DNA is expressed.
protein or polypeptide,
encoded by the
in the cytoplasm or the nucleus of the
However, when it is desired to have the first protein
or polypeptide and/or the third protein or polypeptide
transported from the cytoplasm into chloroplasts or
mitochondria of the cells of transformed plants, the
foreign DNA sequence can further include an additional
foreign DNA encoding a transit peptide. The additional
DNA is between the fertility-restorer DNA and the first
promoter if the first protein or polypeptide is to be
so-transported and is between the first marker DNA and
the second promoter if the third protein or polypeptide
is to be so-transported. By "transit peptide" is meant
a polypeptide fragment which is normally associated
with a chloroplast or mitochondrial protein or subunit
of the protein and is produced in a cell as a precursor
protein encoded by the nuclear DNA of the cell. The
transit peptide is responsible for the translocation
process of the nuclear-encoded chloroplast or
mitochondrial protein or subunit into the chloroplast
or the mitochondria, and during such a process, the
transit peptide is separated or proteolytically removed
from the mitochondrial
chloroplast or protein or
subunit. One or more of such additional DNAS can be
provided in the foreign DNA sequence of this invention
for transporting one or more first or third proteins or
polypeptides as generally described in European patent
publication 0,189,707 and European patent application
(supra) and in: Van den Broeck et al (1985)
Nature 313, 358-363; Schatz (1987) Eur. J. of Bioch.
155, 1-6; and Boutry et al (1987) Nature 328, 340-342.
An example of a suitable transit peptide for transport
into chloroplasts is the transit peptide of the small
subunit of the enzyme RUBP carboxylase (European patent
publication 0,189,707)
peptide for transport into mitochondria is the transit
and an example of a transit
peptide of the enzyme Mn—superoxide dismutase (European
patent publication 0,344,029).
In the foreign DNA sequence of this invention, 3'
transcription termination signals can be selected from
among those which are capable of providing correct
transcription termination and polyadenylation of mRNA
in plant cells. The transcription termination signals
can be the natural ones of the foreign gene or DNA to
be transcribed or can be foreign or heterologous.
Examples of heterologous transcription termination
signals are those of the octopine synthase gene (Gielen
et al (1984) EMBO J. ;, 835-845) and the T-DNA gene?
(Velten and Schell (1985)
("NAR") 1;, 6981-6998).
Nucleic Acids Research
Also in accordance with this invention, a culture
of plant cells, containing the foreign DNA sequence of
this invention, can be used to regenerate homozygous
dominant fertility-restorer plants by performing the
necessary transformation: on a haploid cell culture
(1985) Plant Sci. 3_9_, 219-225)
and then doubling the number’ of chromosomes by well
(Chuong and Beversdof
known techniques (e.g., by the use of colchicine); or
alternatively, on a diploid cell culture and then
culturing anthers of regenerated plants to produce
which can afterwards be rendered
see: Plant Tissue and Cell Culture, Plant
A.R. N.Y. (1987). Thereby, the
foreign DNA sequence will be in homozygous form in the
haploid progeny
diploid.
Biology ;, Liss, Inc.
nuclear genome of each of the so-transformed plant
cells of the culture. This is preferred for a plant
cell culture containing a fertility-restorer DNA under
the control of a first promoter which directs gene
i) the
especially after
expression at a given stage of development of:
plant's male gametes,
the plant's
especially after meiosis, or iii) cells derived
such as pollen,
meiosis, ii) female gametes, such as
ova,
from the male or female gametes, such as seed or embryo
cells, so that the fertility-restorer DNA is present
and can be expressed in all male or female gametes or
plant cells derived therefrom.
with this invention,
processes are provided for producing hybrid seeds which
Further in accordance
can be grown into hybrid fertility-restored plants. one
process involves crossing: a nuclear male-sterile
female-fertile plant including at least one second
marker DNA under the control of at least one third
promoter; with a homozygous nuclear male-fertile
restorer plant including at least one nuclear male
fertility-restorer DNA under the control of at least
one first promoter but without a first marker DNA that
is the same as the second marker DNA. In this process,
the male-sterile and male-fertile plants are sown at
the selectable marker,
random, and after pollination,
encoded by the second marker DNA, is used to eliminate
ensuring that seed is
This
guarantees that all harvested seeds are both hybrid and
the fertility-restorer plants,
only harvested on the male-sterile plants.
fertile. Another process involves crossing: a nuclear
male-sterile female-fertile restorer plant including a
nuclear first marker DNA under the control of a second
promoter and a nuclear female fertility—restorer DNA
under the control of a first promoter in a homozygous
form; with a nuclear male-fertile female-sterile
restorer plant including at least the same nuclear
first marker DNA under the control of a second promoter
and a nuclear male fertility-restorer DNA under the
control of a first promoter in a homozygous form. Both
male-sterile and male-fertile parent plants can be
grown in a substantially random population, thereby
increasing the chances of cross-pollination, without
the need for precise planting patterns, and using the
%
Preferably in
characteristic encoded by the first marker DNA,
fertile hybrid seeds can be harvested.
both of these processes, the first marker DNA is under
the control of a constitutive second promoter and
encodes a third protein or polypeptide that renders the
sterile plant resistant to a particular herbicide. The
non—desirab1e genotypes can then be destroyed before
cross-pollination, using the particular herbicide.
A. process in accordance with this invention of
1) fertility-restorer plants which contain a
—DNA stably
nuclear genome and transmissible throughout generations
crossing:
fertility—restorer integrated in their
as a dominant allele in accordance with this invention,
with 2) male- or female-sterile plants which contain a
sterility DNA,
second marker DNA,
preferably both a sterility DNA and a
stably integrated in their nuclear
genome and transmissible throughout generations as
dominant alleles in accordance with European patent
0,344,029 and 0,412,006,
and several advantages over,
publication provides an
alternative to, presently
used systems for breeding and producing hybrid crops as
described below:
. For crops which do not easily cross-pollinate and
for which the seed is the economic harvest and has
low multiplication rates, such as cereals (e.g.,
wheat, barley and oats), rice, cotton, and many
legumes (e.g., soybean and pea), the process of
this invention offers the possibility to produce
100% hybrid fertile offspring, thereby
guaranteeing high seed set and normal yield. An
example of a typical strategy for producing hybrid
plants, using as parent plants male-sterile and
female-sterile parent plants and a restorer for
their respective sterilities, may include the
following steps (wherein "FH1" stands for female-
"RF"
stands for the restorer of the female—sterility,
"M1H1" linked to
herbicide resistance 1, stands for male-
sterility linked to herbicide resistance 1,
stands for male-sterility l
”M2H2"
sterility 2 linked to herbicide resistance 2,
"RM1" stands for restorer of male—sterility 1, "A"
stands for female parent lines, and "B" stands for
male parent lines):
A. Development of the female parent plant A
1Aa) Transform plant A with a fertility-restorer
DNA of this invention that encodes a first
RNA, (which
neutralizes expression
protein or polypeptide
specifically the
lAc)
lAd)
product of the female—sterility’ DNA in the
male parent) and is under the control of a
first promoter which directs gene expression
in at least the same cells as those in which
the female-sterility DNA in the male plant is
to be expressed. This gives rise to ARF/rf.
Self-pollinate ARF/rf, giving rise to 25 %
ARF/RF plants.
ARF/RF with a
sequence including a "male-sterility DNA 1"
Transform chimaeric DNA
under the control of a male organ specific
promoter and a marker DNA conferring
resistance to a herbicide 1. This gives rise
to the male-sterile plant ARF/RF7M1H1/mh.
Multiply the male-sterile plant by crossing:
ARF/RF:MlH1/mh X ARF/RF:mh/mh
giving an offspring consisting of:
% ARF/RF’M1H1/mh: male-sterile 1, resistant
to herbicide 1 and
% ARF/RF7mh/mh: male fertile, herbicide
sensitive.
This mixture is sown in successive
generations of upscaling of the female
parent, and the herbicide 1 is used in
alternate rows or blocks of rows to create
pure female parent stocks. The rows or blocks
of rows where the herbicide is not applied
are used as pollen source. only seed of the
herbicide treated rows or blocks of rows is
harvested to constitute the next generation.
B. Development of the male parent plant B
For the economic production of B, the female-
sterile parent line requires the use of two
different sterility DNAs. The first one is a
female-sterility DNA under the control of a
promoter which directs gene expression selectively
in cells of the female organ of the plant and
linked to a marker DNA conferring resistance to
herbicide l. The second one is a male-sterility
DNA (different from male—sterility DNA 1 and
called "male-sterility DNA 2"), under the control
of a promoter which directs gene expression
selectively in male organ cells of the plant and
is linked to a second marker DNA conferring
resistance to herbicide 2.
1Ba) Transform plant B with a foreign DNA sequence
of this invention encoding a first RNA,
protein or polypeptide which neutralizes
specifically the activity of the male-
sterility DNA 1 expressed in the female
parent and is under the control of a first
promoter which directs gene expression in at
least the same male organ cells in which the
male-sterility DNA 1 in the female parent
plant is expressed. This gives rise to
BRMI/rm_
lBb) Self-pollinate BRM1/rm, giving rise to 25%
BRMl/RMl_
1Bc) (1) Transform BRM1/RM1 with a chimaeric DNA
sequence including the female-sterility DNA
under the control of a promoter which directs
gene expression selectively in cells of the
female-organ of the plant and a marker DNA
lBd)
conferring resistance to herbicide 1. This
gives rise to the
BRM1/RM1:FHl/fh_
female-sterile plant
(2) Transform BRM1/RM1 with a chimaeric DNA
sequence including the male-sterility DNA 2
under the control of a male organ-specific
promoter and a marker DNA conferring
herbicide resistance to herbicide 2. This
gives rise to the
BRMI/RMl;M2H2/mh_
male-sterile plant
(1) Multiply the male-sterile plant of
1Bc)(2) by crossing:
BRMI/RM1:M2H2/mh X BRMl/RMl;mh/mh
giving an offspring consisting of:
% BRM1/RM17M2H2/mh: male-sterile,
resistant to herbicide and
% BRM1/RM17mh/mh: male fertile, herbicide
sensitive.
(2) Multiply the female-sterile plant of
1Bc)(1) by crossing:
BRM1/RM1;M2H2/mh;fh/fh X
BRM1/RM1;mh/mh7FH1/fh which are planted in
separate rows, giving rise to the following
genotypes in the male-sterile rows:
% BRMI/RM1:M2H2/mh;FHl/fh: sterile and
resistant to herbicide 1 and 2,
% BRM1/RM17mh/mh7FH1/fh: female-sterile
and resistant to herbicide 1,
% BRM1/RM17M2H2/mh7fh/fh: male-sterile and
resistant to herbicide 2, and
% BRM1/R“17mh/mh7fh/fh: fertile and
herbicide sensitive.
Be) This mixture can be used again as the male
parent (BRM1/RM17mh/mh7FH1/fh) in further
multiplication crosses, whereby spraying in
each generation with herbicide 1 eliminates
the female fertile plants and so maintains
the male parent line. This mixture will be
planted in alternate rows or blocks of rows
with the mixture obtained in 1 Bd (1), which
mixture will be treated with herbicide 2 to
eliminate male fertile plants. Alternatively,
the mixture obtained in 1 Bd) (2) can be sown
as such and alternate rows can be treated
either with herbicide 1 or either with
herbicide 2. Under such circumstances, step 1
Bd)(1) is not necessary.
C. Production of hybrid seed AB
Sow at random the mixtures obtained in the steps
1Ad) and 1Be). Before cross—pollination occurs,
spray with herbicide 1 in order to eliminate all
undesirable genotypes. Cross pollination occurs
with:
ARF/RF; rm/rm;M1Hl/mhzfh/fh X BRI‘/Il/RM1; rf/rf; mh/mh;FH1/fh,
giving rise to:
% ABRF/rf;M1H1/mh;rm/RM1;FH1/fh
% ABRF/rf;M1H1/mh;rm/RM1;fh/fh
% ABRF/rf;mh/mhirm/RM1:FH1/fh
% ABRF/rf;mh/mh;rm/RM1:fh/fh
constituting 100 % fertile hybrid seed.
Depending on ‘the special characteristics of the
crop which is bred, the foregoing general strategy
can be simplified. Such special characteristics
include:
(2.1)
(2.2)
If the crop undergoes a reasonable or good
cross-pollination by insects, the relative
proportion of parent line B in the mixture
can be lowered without affecting the yield of
the crop (e.g., cotton, a legume such as
Pisum, alfalfa, oilseed rape and corn).
Alternatively, a much simplified breeding
scheme can be used for" a crop involving a
female parent which has been rendered male-
sterile and herbicide resistant and. a male
parent carrying the fertility—restorer' gene
for the male-sterility.
This would permit the following strategy:
Cross: AMH/mh x BR”/RM sown at random or in
rows for crops which do not flower
synchronously.
Treat with herbicide after pollination when
sown at random.
Yielding: so % ABMH/mh7RM/rm and so % ABmh/mh7RM/rm,
constituting 100 % fertile hybrid offspring.
In case F2 offspring represent the commercial
(e.g.
variant strategy can be used:
seed product cotton), the following
Produce by transformation male—sterile plants
of parent line A, giving AM/m’r/r:
transformation
Produce by 2 independent
events fertility-restorer plants carrying
into two independent genetic loci of its
nuclear genome the fertility-restorer gene
the product of which neutralizes specifically
the activity the male-sterility gene in the
male—steri1e plant of a) and obtain by self-
pollination
both
restorer
genes
homozygous form, giving Bm/m7R1/R17R2/R2;
C) Cross AM/m:r/r X Bm/m;R1/Rl:R2/R2
yielding 50 % ABM/m7R1/r7R2/r and 50 %
ABm/m;R1/r7R2/r
constituting 100 % hybrid fertile offspring;
and
d) self-pollinate the mixture obtained in c).
Half of the offspring are as shown in Table
1, below, only 1 of a total of 64 plants
being male-sterile (indicated by an * in
Table 1), and all the others being fertile.
This result makes this process economically
valuable.
Table 1
AB\AB sRlR2 sR1r2 srlR2 sr1r2
SR1R2 SRlR2/SR1R2 sR1r2/SR1R2 srlR2/SR1R2 sr1r2/SR1R2
SRlr2 sR1R2/SrlR2 SRlr2/SRlr2 srlR2/SRlr2 sr1r2/SRlr2
srlR2 sRlR2/srlR2 sR1r2/srlR2 srlR2/srlR2 sr1r2/Sr1R2
Sr1r2 sR1R2/sr1r2 sRlr2/Srlrz srlR2/Sr1r2 sr1r2/Sr1r2*
sR1R2 sR1R2/sRlR2 sR1r2/SR1R2 srlR2/sR1R2 sr1r2/sR1R2
SRlr2 sR1R2/sR1r2 sR1r2/sR1r2 srlR2/SRlr2 sr1r2/SRlr2
srlR2 sR1R2/srlR2 sR1r2/srlR2 srlR2/srlR2 sr1r2/srlR2
sr1r2 sR1R2/sr1r2 sR1r2/srlrz srlR2/srlrz sr1r2/sr1r2
in a
(2.3)
(2.4)
(2.5)
A.
3Aa)
If the male-sterility DNA 2 is linked to
another marker DNA than the one encoding
resistance to herbicide 2, e.g. a color gene,
the plants carrying this male-sterility DNA
could be easily eliminated without damage to
the other plants. Alternatively, the male-
sterility DNA 2 could be introduced without
any selectable marker DNA. Eliminating plants
carrying the male—sterility DNA 2 could be
done through manual selection, which needs
only to be done on a small scale (See (1)
Bd), above).
If the tissue of the parent plants to be
transformed is constituted of haploid
material, this would reduce considerably the
subsequent breeding, presenting the dominant
genes encoding sterility in a homozygous
form.
If the value of the seed, or the cost of hand
labor allows manual elimination of unwanted
genotypes, at least up to the last stages
before the hybrid production, the general
system could also be simplified.
Another example of a breeding strategy-—using
male- and female-sterility combined with the
fertility
invention--may include the following steps:
restorer system of this
Development of the female parent line A
Transform line A with a foreign DNA sequence
including a fertility-restorer DNA of this
invention which: encodes a first RNA, protein
or polypeptide that neutralizes specifically
Ac)
Ad)
the activity of the product of a female-
sterility DNA expressed in the male parent;
is under the control of a first promoter that
directs expression of the fertility-restorer
DNA in at least the same female organ cells
as those in which the female—steri1ity DNA of
the male parent is expressed; and is adjacent
to a first marker DNA encoding resistance to
herbicide 2. This gives rise to ARFH2/rfh.
Transform also, in parallel, line A with a
DNA sequence including a male-sterility DNA
which: is under the control of a male organ-
specific promoter; and is adjacent to a
second marker DNA encoding a different
herbicide resistance (i.e., to herbicide 1)
from the one encoded by the first marker DNA.
This gives rise to AMH1/mh
Cross ARFH2/rfh X AM1/mh,
% ARFH2/rfh:MH1/mh
% ARFH2/rfh;mh/mh
% Arfh/rfh;MH1/mh
% Arfh/rfh:mh/mh_
giving rise to
Spray with herbicides 1 and 2, selecting
ARFH2/rfh;M1/mh_
Self-pollinate ARFH2/rfh X ARFH2/rfh,
giving rise to 25% ARFH2/RFH2, which can be
maintained by self-pollination.
Cross ARFH2/RFH2;mh/mh X ARFH2/rfh: MR1/mh_
This gives rise to:
% ARFH2/RFH2; MH1/mh
% ARFH2/RFH2: mh/mh
% ARFH2/rfh; Ml/mh
Ae)
Ba)
% ARFH2/rfh; mh/mh
whereby the male—steri1e plants, having the
fertility-restorer DNA in homozygous form,
can be selected by spraying with herbicide 1
and by test-crossing with,parenta1 A line.
female
Maintain the parent line A by
crossing:
ARFH2/RFH2;MH1/mh X ARFH2/RFH2:mh/mh_
Development of the male parent line B
Transform line B with a foreign DNA sequence
V including a fertility-restorer DNA. of this
invention which: encodes a first RNA, protein
or polypeptide that neutralizes specifically
the activity of the product of a male-
'sterility DNA expressed in the female parent;
is under the control of a first promoter that
directs expression of the fertility-restorer
DNA in at least the same male organ cells as
those in which the male—steri1ity DNA is
expressed; and is adjacent to a first marker
DNA encoding resistance to herbicide 2. This
gives rise to BRMH2/rmh.
Transform, in parallel, also line B with a
DNA sequence including a female-sterility DNA
which: is under the control of a female
organ—specific promoter; and is adjacent to a
second marker DNA encoding resistance to
herbicide 1. This gives rise to BFH1/fh.
Cross BRMH2/rmh:fh/fh X Brmh/rmh:FH1/fh,
giving rise to:
% BRM2/rmh; Ffll/fh
% BRMH2/rmh’ f9/E3
Bc)
Bd)
Be)
Ca)
Isolate BRM2/rmh7FH1/fh by spraying with
herbicides 1 and 2.
Self-pollinate BRMH2/rmh X BRKHZ/rm“,
giving rise to:
% BRM2/RMH2
which can be maintained
through self-pollination.
Cross BRMH2/RMM2 X BRMBZ/rmh:FH1/fh
giving rise to:
% BRMH2/RMH2:FH1/fh
% BRMH2/RMH2;fh/fh
% BRMH2/rmh:FH1/fh
% BRMH2/rmh:fh/fh
whereby the female-sterile plants having the
fertility-restorer DNA in homozygous form are
selected by spraying with herbicide 1 and by
test-crossing with parental B line.
Maintain the male parent line B by crossing:
BRMH2/RM2:fh/fh X BRM2/RMH2;FH1/fh_
Alternative procedure for development of male
or female parent plant (A or B are both
designated by C)
Transform line C with a foreign DNA sequence
including a fertility-restorer DNA of this
invention which: encodes a first RNA, protein
or polypeptide that neutralizes specifically
the activity of the product of a sterility
DNA expressed in the other parent; is under
the control of a first promoter that directs
expression of the fertility restorer DNA in
at least the cells in which the sterility DNA
Cb)
Cc)
Cd)
of the other parent is expressed: and is
adjacent to a first marker DNA encoding
resistance to herbicide 2. This gives rise to
CRH2/rh_
Self-pollinate CRH2/rh X CRH2/rh, producing
% CRH2/RH2 which can be maintained through
self-pollination.
CRH2/RH2 with a DNA
including a sterility DNA which is: under the
Transform sequence
control of a male or female organ-specific
promoter and adjacent to a second marker DNA
encoding resistance to herbicide 1. This
gives rise to CRH2/RH27sH1/sh (wherein "S"
stands for male- or female-sterility).
Maintain line C by the following cross:
CRHZ/RH2:SH1/sh X CRH2/RH2;sh/sh_
Production of hybrid seed AB
Sow at random the mixtures obtained in steps
3Ae) and 3Be) or the mixture obtained in step
3Cd). Before cross-pollination occurs, spray
with herbicides 1 and 2 in order to eliminate
all undesirable genotypes. This leads to the
following cross:
ARFH2/RFH2;rm/rm;MH1/mh;fh/fh X
BRMH2/RMH2;rf/rf;FH1/fh;mh/mh_
This gives rise to the following offspring:
% ABRFH2/rf;RM2/rm;Mm1/mh:FH1/fh
% ABRFH2/rf:RM2/rm;MH1/mh;fh/fh
% ABRFH2/rf;RMH2/rm;mh/mh;FH1/fh
% ABRFH2/rf;RMm2/rm;mh/mh;fh/fh
consisting of 100% hybrid fertile seed.
. other advantages of the fertility-restorer
system of this invention, combined with the
male- or female-sterility systems described
in European patent publications 0,344,029 and
,412,006, compared to earlier systems,
include:
a) A fool-proof production scheme, with several
well distinguishable and selectable markers
to control quality;
b) A considerable reduction in complexity at the
level of the final seed multiplier, which is
essential for reliable production and reduced
production costs; and
c) Reduction of the time necessary for the
production of a commercial hybrid seed.
The following Examples illustrate the invention.
The figures referred to in the Examples are as follows:
Fig. 1 shows a map of the vector pTVE74 of
Example 1.
Fig. 2 shows the DNA sequence of the barstar
and indicates the
gene, used. in Example 1,
mutated sequence of its ClaI site.
Unless otherwise stated in the Examples, all procedures
for making and manipulating recombinant DNA were
carried out by the standardized procedures described in
Maniatis et al, Molecular Cloning - A Laboratory
Manual, Cold Spring Harbor Laboratory (1982). The
following plasmids and vectors, used in the Examples,
Deutsche Sammlung Fur
("DSH').
Federal
have been deposited in the
Mikroorganismen und zellculturen
Weg 1B, D-3300
Mascheroder
Republic of
Germany under the provisions of the Budapest Treaty:
Braunschweig,
Plasmid DSM Accession
or No. Date
vector
pMB3 4470 21 Mar. 1988
pGSCl70O 4469 21 Mar. 1988
Example 1 - Construction of a chimaeric DNA sequence
of PTA29 and a barstar gene
A plasmid named "pTVE74", shown in Fig. 1, is
constructed by assembling the following well known DNA
fragments with the PTA29 promoter:
. a vector fragment, including T-DNA border
sequences, derived from pGSC1700 (cornelissen and
Vandewiele (1989) NAR 11 (1) 19-29) in which the
B-lactamase gene has been deleted; located between
the border sequences are the following DNA
fragments 2 and 3;
. a chimaeric sequence containing the promoter
cassette PTA29 from European patent publication
,344,029, fused in frame at the ATG initiation
codon with a Bacillus amyloliquefaciens gene
encoding barstar, which is the cellular inhibitor
of the
(Hartley et al (1972) Preparative Biochemistry g
extracellular ribonuclease, Barnase
) 243-250; Hartley and Smeaton (1973) J. Biol.
Chem. 248 (16), 5624»5626); the following" steps
are carried out:
a) The nucleotide sequence GCAC, at positions 7
to l0 up—stream of the first ATG codon, is
mutated into nucleotide sequence ATCG, in
order to obtain a suitable Clal cloning site
at the first methionine codon of the coding
sequence (see Fig.2); this is accomplished
using site directed mutagenesis (European
patent application 87402348.4 which is the
priority application claimed in European
patent publication 0,319,353) and yields
pMc5—TPBSC; the Clal protruding ends are
digested by the enzyme, SI, and the barstar
gene is isolated as a ClaI—HindIII fragment
of 330 nucleotides (Fig. 2); and
b) The SI—treated ClaI—HindIII fragment of pMc5—
TPBSC is fused with the SI—treated NcoI—
Hindlll fragment of pMB3 (European Patent
publication 0,344,029) and with a restriction
fragment containing the 3' end signals of the
nopaline synthetase ("NOS") gene for
transcription termination and polyadenylation
(An et al (1985) EMBO J. 4 (2), 277); and
. a chimaeric sequence containing an Arabidopsis
Rubisco SSU promoter ("PSSU" or "PSSUARA"), a neo
gene encoding, kanamycin resistance (European
patent publication 0,242,246) and the 3' end
signals of the octopine synthase ("OCS”) gene
(Dhaese et al (1983) EMBO J. 2, 419).
pTVE74 is a binary type T—DNA vector containing,
within the T~DNA border sequences, three chimaeric
sequences: PNOS-neo and PSSU-sfr which comprise first
marker DNAS under the control of their own second
promoters: and PTA29-barstar in which barstar is a
fertility-restorer DNA whose expression under the
control of the tapetum-specific PTA29 first promoter
will neutralize, in tapetum cells of an otherwise
male-sterile plant, the activity of Barnase encoded by
a sterility DNA under the control of a tapetum-specific
sterility promoter as described in European patent
publication 0,344,029.
of the
sequence of Example 1 into tobacco and oilseed rape
Example 2 - Introduction chimaeric DNA
A recombinant Agrobacterium strain is constructed
by mobilizing pTVE74 (from Example 1) from E. coli into
Agrobacterium tumefaciens CSBC1 RifR containing pMP90
(Koncz and Schell (1986) Mol. Gen. Genetics 204,
-396). The resulting Agrobacterium strain harboring
pMP90 and pTVE74 is used to transform tobacco leaf
discs (Q. tabacum Petite Havane SR1) using standard
procedures as described, for example, in European
patent publication 0,242,246 and to transform oilseed
according to the procedure of
, 464-466 and
rape (Brassica napus)
Lloyd et al (1986)
(1985)
Carbenicillin is used to kill the
Science
Klimaszewska et al Plant Cell Tissue Organ
Culture 4, 183-197.
Agrobacterium strains after infection.
Transformed calli are selected on substrate
containing 100 ug/ml kanamycin, and resistant calli are
regenerated into plants. After induction of shoots and
transferred to the
roots, the transformants are
greenhouse and grown until they flower. The flowers are
examined, and they exhibit a fully natural morphology.
Pollens of these flowers are used to pollinate the
nuclear male-sterile tobacco and oilseed rape plants
containing the Barnase gene as a sterility DNA, under
the PTA29
sterility promoter, described in Example 13 of European
the control of tapetum cell—specific
patent publication 0,344,029. Offspring of these
pollinated male-sterile plants are analyzed, and 75% of
their flowers do not exhibit a male-sterility phenotype
(i.e., absence of a normal tapetum layer in the stamens
of their flowers).
Needless to say, this invention is not limited
to the transformation of any specific plant(s). The
invention relates to any plant, the nuclear genome
fertility-
restorer DNA under the control of a first promoter
that can of the fertility-
restorer DNA selectively in at least cells of the
of which can be transformed with a
direct expression
plant's flowers, particularly at least one male or
at least one female organ thereof, and/or seeds
and/or embryos, whereby the plant can be both
self-pollinated and crdss—po1linated.
this
For example,
invention relates to plants such as corn,
oilseed rape, wheat, rice, sunflower, sugarbeet,
tomato,
alfalfa,
lettuce, peppers, sorghum,
clovers,
soybean, pea,
leek,
onion, tobacco, petunia, cacao and citrus trees.
Also,
grasses, carrot, cabbages,
this invention is not limited to the
specific plasmids and vectors described in the
foregoing Examples, but rather encompasses any
plasmids and vectors containing the fertility-
restorer DNA under the control of the first
promoter.
Furthermore, this invention is not limited to
the specific first promoters described in the
foregoing Examples, such as the PTA29 promoter, but
rather encompasses any DNA sequence encoding a
first promoter capable of directing expression of a
fertility-restorer DNA at least in cells of a
plant's flowers, seeds and/or embryos, where
expression of a sterility DNA would otherwise cause
the plant to be male— or female-sterile. In this
regard, the first promoter of this invention
encompasses: the promoters described in European
patent publication 0,344,029 for use in
controlling the expression of a sterility DNA
selectively in stamen cells of a plant to be
rendered male-sterile; and the promoters described
in European patent publication 0,412,006 for use
in controlling the expression of a sterility DNA
selectively in cells of flowers, seeds or embryos
of a plant to be rendered female-sterile.
Alternatively, the first promoter can be a
constitutive promoter for the plant, provided the
first RNA, protein or polypeptide does not
significantly disturb adversely the functioning,
metabolism or development of cells in which it is
expressed in the absence of expression of the
sterility DNA.
In addition, this invention is not limited to
the specific fertility-restorer DNAs described in
the foregoing Examples but rather encompasses any
DNA sequence encoding a first protein or
polypeptide which, in a fertility-restored plant,
neutralizes, blocks, offsets, overcomes or
otherwise prevents the activity of the second protein or
polypeptide which is a ribonuclease that is encoded by the
sterility-DNA under the control of the sterility
promoter and that would otherwise significantly
disturb adversely the metabolism, functioning
and/or development of cells of flowers, seeds or
embryos of the plant.
Also, this invention is not limited to the
specific first marker DNAs described in the
foregoing Examples but rather encompasses any DNA
sequence encoding a third RNA, protein or
polypeptide which confers on at least a specific
plant tissue or specific plant cells, in which such
DNA sequence is expressed, a distinctive trait
compared to such a specific plant tissue or
specific plant cells in which such DNA sequence is
not expressed.
Claims (6)
1 A recombinant DNA comprising a first chimeric DNA which comprises : at a restorer DNA encoding a protein which is an inhibitor of a ribonuclease. and, b) a first promoter which directs expression at least in specific cells of a flower, a seed and/or an embryo of a plant. and wherein said restorer DNA is in the same transcriptional unit as. and under the control of, said first promoter.
2 The recombinant DNA of claim‘ 1 in which said inhibitor is capable of neutralizing the activity of the extracellular ribonuclease barnase of Bacillus amvloliouefaciens.
3. The recombinant DNA of claim 2in which said inhibitor is barstar with an amino acid sequence as encoded by the coding sequence starting at nucleotide position 11 in Figure 2.
4. The recombinant DNA of claim 3ln which said restorer DNA comprises the coding sequence starting at nucleotide position 11 in Figure 2.
5. The recombinant DNA of claim 4 in which said restorer DNA is the Clal-Hindlll fragment of Fig. 2.
6. The recombinant DNA of any one of claims 1 or 5, which also comprises a second chimeric DNA, comprising: comprising : (c) a marker DNA encoding a marker RNA. protein or polypeptide which, when present at least in a specific tissue or in at least specific cells of a plant, renders said plant easily separable from other plants which do not contain said marker RNA, protein or polypeptide in said specific tissue or specific cells; and (d) a second promoter capable of directing expression of said marker DNA at least in said specific tissue or specific cells; said marker DNA being in the same transcriptional unit as, and under the control of, said second promoter. The recombinant DNA of claim 6 in which said marker DNA encodes a protein or polypeptide conferring a color to at least said specific tissue or specific cells; or encodes a protein or polypeptide conferring on said plant a stress tolerance. a disease or pest resistance or a bacterial resistance. The recombinant DNA of claim 7 wherein said marker DNA encodes a Bacillus thuringiensis endotoxin that centers insect resistance, or encodes a bactericidal peptide that confers a bacterial resistance. The recombinant DNA of claim 6 wherein said marker DNA encodes a modified target enzyme for a herbicide having lower affinity for the herbicide than the unmodified target enzyme. The recombinant DNA of claim 9 wherein said marker DNA encodes a protein or polypeptide which is selected from the group of a modified 5-enolpyruvylshikimate-3 phosphate synthase as a target for the herbicide. glyphosate and a modified glutamine synthetase as a target for a glutamine synthetase inhibitor. including phosphinothricin. The recombinant DNA of claim 6 wherein said marker DNA encodes a protein or polypeptide that inhibits or neutralizes the activity of a herbicide. ' The recombinant DNA of claim 11 wherein said marker DNA encodes a protein or polypeptide S0 conferring resistance to a glutamine synthetase inhibitor, including phosphinothricin. The recombinant DNA of claim 12 wherein said marker DNA is _a sir or sfrv gene. The recombinant DNA of any one or’ claims 6 to 13 wherein said second promoter is a constitutive promoter, a wound-inducible promoter, a promoter which directs expression selectively in plant tissue having photosynthetic activity, or a promoter which directs gene expression selectively in leaf cells. petal cells or seed cells. The recombinant DNA of claim 14 wherein said second promoter is a 35S promoter, a 35S'3 promoter, a Pnos promoter, a TR1‘ or TR2’ promoter, or a SSU promoter. The recombinant DNA of any one of claims 1 to 15 which also comprises: (e) a first DNA encoding a transit peptide capable of transporting said inhibitor into a chloroplast or mitochondria of said stamen cells: said first DNA being in the same transcriptional unit as said restorer DNA and said first promoter and between said restorer DNA and said first promoter: and/or (f) a second DNA encoding a transit peptide capable of transporting said marker protein or polypeptide into a chloroplast or mitochondria of at least said specific tissue or specific cells; said second DNA being in the same transcriptional unit as said marker DNA and said second promoter and between said marker DNA and said second promoter. The recombinant DNA of any one of claims 1 to 16 in which said first promoter is a constitutive promoter. The recombinant DNA’rof‘any‘one‘ofElaimis'1 to 16 in which said first promoter directs expression at least in cells of the male organ of a plant. The recombinant DNA of claim 18 in which said first promoter directs expression selectively in stamen cells of a plant. 5 1 The recombinant DNA of claims 18 or 19 in which said first promoter directs expression in one or more types of stamen cells selected from the group The recombinant DNA of claim 20 in which said first promoter directs expression in anther cells. The recombinant DNA of claim 21 in which said first promoter is a promoter from an endogenous plant gene selected from the group of the TA29 gene from tobacco, the TA26 gene from tobacco, the TA13 gene from tobacco, a gene encoding a mRNA hybridizable to said TA29 gene, a gene encoding a mRNA hybridizable to said TA13 gene, and a gene encoding a mRNA hybridizable to said TA25 gene. The recombinant DNA of claim 22 in which said first promoter is the TA29 promoter contained in the Ncol-Hindlll fragment of plasmid pMB3, DSM 4470. The recombinant DNA of any one of claims 1 to 16 in which said first promoter directs expression at least in cells of the female organ of a plant. The recombinant DNA of claim 24 in which said first promoter directs expression selectively in cells of the female organs of a plant. The recombinant DNA of claims 24 or 25 in which said first promoter directs expression in one or more types of cells of the lemale organ selected from the group of ovary, ovule, style, stigma, or septum cells. The recombinant DNA of..c|aim-26_in_which said first promoter is a promoter that directs expression in style and/or stigma cells, such as the promoter of the endogenous STGM4B12 gene of tobacco, the endogenous STGM3C9 gene of tobacco, the promoter of the endogenous STGM07 gene from tobacco, or the promoter of the endogenous .STGM08 gene from tobacco. The recombinant DNA of any one of claims 1 to 27 which is part of the nuclear genome of a cell of a plant or of a seed. of anther, pollen. and filament cefls. The recombinant DNA ol claim 28 which also contains a DNA encoding said ribonuclease. A cell of a plant which contains the recombinant DNA of any one of claims 1 to 29. A cell of claim 30 which can be regenerated into a plant. A plant which contains the DNA of any one of claims 1 to 29. A plant which contains the recombinant DNA of any one of claims 1 to 29 in all of its cells. A seed of a plant wherein said seed contains the recombinant DNA of any one of claims 1 to 29. The seed of claim 34 which is a hybrid seed. A plant which contains integrated into the nuclear DNA or’ all of its cells. the recombinant DNA of any one oi claims 17 to 23 and which is capable. when crossed to a second plant which contains a second recombinant DNA comprising a sterility DNA encoding said ribonuclease under the control of a sterility promoter which directs exoression selectively in specific stamen cells of said second plant and which is male-sterile due to the selective production of said ribonuclease in said specific stamen cells. of producing male fertile progeny plants that produce said ribonuclease and said inhibitor of said ribonuclease in said specific stamen cells. The plant of claim 36 which is homozygous tor said recombinant DNA. A plant which contains integrated into the nuclear DNA of all of its cells. a) the recombinant DNA of any one of claims 17 to 23. and. b) a second recombinant DNA comprising a sterility DNA encoding said ribonuclease under control of a sterility promoter which directs expression selectively in specific 53 stamen cells of said plant. and which is male fertile due to the neutralizing of the activity of said ribonuclease in said specific stamen cells by said inhibitor produced by expression of said restorer DNA at least in said specific stamen cells. The plant of claim 38 which is a hybrid plant. The plant of any one of claims 36 to 39 in which said ribonuclease is a barnase and said restorer DNA encodes barstar. The plant of any one of claim 36 to 40 in which said sterility promoter directs expression in one or more types of stamen cells seiected from the group of anther. pollen. and filament cells. The plant of claim 41 in which said sterility promoter directs expression in one or more types of stamen cells selected from the group of tapetum and anther epidennal cells. The plant of claim 42 in which said sterility promoter is a promoter from an endogenous plant gene selected from the group of the TA29 gene from tobacco. the TA26 gene from tobacco. the TA13 gene from tobacco. a gene encoding a mRNA hybridizable to said TA29 gene, a gene encoding a mRNA hybridizable to said TAl3 gene. and a a gene encoding a mRNA hybridizable to said TA26 gene. The plant of claim 43 in which said sterility promoter is the TA29 promoter contained in the Ncol- Hindlll fragment of plasmid pMB3, DSM 4470. The plant of any one of claims 36 to 44 in which said first promoter and said sterility promoter are the same. A pair of parent plants for producing seeds comprising: (a) a male-sterile parent plant which contains incorporated in the nuclear genome of all of its cells, a sterility DNA encoding said ribonuclease under the control of a sterility promoter which directs expression of said sterility DNA selectively in specific stamen cells of said plant. and. (b) a male-fertile parent plant which contains incorporated into the nuclear DNA of all of its cells. the recombinant DNA of any one of claims 17 to 23 wherein said first promoter directs expression at least in the same specific stamencells as said sterility promoter: and whereby said ma|e—sterile and male-fertile plant can be crossed to produce male-fertile progeny comprising said recombinant DNA and said sterility DNA under control of said sterility promoter. The pair of claim 46 in which said ribonuclease is bamase and said restorer DNA encodes barstar The pair or’ claim 46 or 47 in which said sterility promoter directs expression in one or more types of stamen cells selected from the group of anther. pollen, and filament cells. The pair of claim 48 in which said sterility promoter directs expression in one or more types of stamen cells selected from the group of tapetum and anther epidermal cells. The pair of claim 49 in which said sterility promoter is a promoter from an endogenous plant gene selected from the group oi the TA29 gene from tobacco, the TA26 gene from tobacco. the TA13 gene from tobacco. a gene encoding a mRNA hybridizable to said TA29 gene, a gene encoding a mFlNA hybridizable to said TA13 gene. and a a gene encoding a mRNA hybridizable to said TA26 gene. The pair of claim 50 in which said sterility promoter is the TA29 promoter contained in the Neol- Hindlll fragment of plasmid pMB3. DSM 4470. The pair of any one of claims 46 to 51 in which said first promoter and said sterility promoter are the samef The pair of any one of claims 46 to 52in which said male-fertile parent plant is homozygous for said recombinant DNA. The pair of any one of claims 46 to 53 for producing hybrid seeds. A process for producing a transgenic plant, or reproduction material or progeny plants thereof, which comprises the steps of: a) transforming a starting cell of a plant with the recombinant DNA of any one of claims 1 to 27 to produce a transformed plant cell which contains said recombinant DNA stably integrated into its nuclear DNA: b) regenerating said transgenic plant from said transformed cell. and optionally, cl propagating" said transgenic plant to obtain said reproduction material or progeny plants which contain said recombinant DNA. A plant which contains integrated into the nuclear DNA of all of its cells, the recombinant DNA of claim 17 or of any one of claims 24 to 27 and which is capable, when crossed to a second plant which contains a second recombinant DNA comprising a sterility DNA encoding said ribonuclease under control of a sterility promoter which directs expression selectively in specific cells of the female organ of said second plant and which is female- sterile due to the selective production of said ribonuclease in said specific cells, of producing female fertile progeny plants that produce said ribonuclease and said inhibitor of said ribonuclease in said specific cells. The plant of claim 56 which is homozygous for said recombinant DNA. A plant which contains integrated into the nuclear DNA of all of its cells, a) the recombinant DNA of claim 17 or of any one of claims 24 to 27, and, b) a second recombinant DNA comprising a sterility DNA encoding said ribonuclease under control of a sterility promoter which directs expression selectively in specific cells of the female organ of said plant, and which is female fertile due to the neutralizing of the activity of said ribonuclease insaid specific cells by said inhibitor produced by expression of said restorer DNA at least in said specific cells. The plant of claim 58 which is a hybrid plant. The plant of any one of claims 56 to 59 in which said ribonuclease is bamase and said restorer DNA encodes barstar. The plant of any one of claim 56 lo 60 in which said sterility promoter directs expression in one or more types of cells selected from the group of ovary, ovule. style. stigma and septum cells. The plant of claim 61 in which said first promoter is a promoter that directs expression in style and/or stigma cells. such as the promoter of the endogenous STGM4B12 gene of tobacco. the endogenous STGMBCQ gene of tobacco. the promoter of the endogenous STGMD7 gene from tobacco, or the promoter of the endogenous STGM08 gene from tobacco. The plant of any one of claims 56 to 62 in which said first promoter and said sterility promoter are the same. A pair of parent plants for producing seeds comprising: (a) a female-sterile parent plant which contains incorporated in the nuclear genome of all of its cells. .3 sterility DNA encoding said ribonuclease under the control of a sterility promoter which directs expression of said sterility DNA selectively in specific cells of the female organ of said plant, and. (b) a female-fertile parent plant which contains incorporated into the nuclear DNA of all of its cells, the recombinant DNA of claim 17 or of any one of claims 24 to 27 wherein said first promoter directs expression at least in the same specific cells of the female organ as said sterility promoter; and whereby said fema|e—sterile and female—terti|e plant can be crossed to produce female-fertile progeny comprising said recombinant DNA and said sterility DNA under control of said sterility promoter. The pair of claim 64 in which said ribonuclease is bamase and said restorer DNA encodes barstar. The pair of claim 64 or 65 in which said sterility promoter directs expression in one or more types of cells selected from the group of ovary, ovule, style, stigma and septum cells. The pair of claim 66 in which said sterility promoter is a promoter that directs expression in style and/or stigma cells, such as the promoter of the endogenous STGM4B12 gene of tobacco, the endogenous STGM3C9 gene of tobacco, the promoter of the endogenous STGM07 gene from tobacco, or the promoter of the endogenous STGM08 gene from tobacco. The pair of any one of claims 64 to 67 in which said first promoter and said sterility promoter are the same. The pair of any one of claims 64 to 68 in which said female-fertile parent plant is homozygous for said recombinant DNA. The pair of any one of claims 64 to 69 for producing hybrid seeds. The use of the recombinant DNA of any one of claims 1 to 28 to produce an inhibitor of a ribonuclease in at least specific cells of a flower, a seed and/or an embryo of a plant. The use of claim 71 to neutralize the activity of said ribonuclease produced in said at least specific cells. F. R. KELLY & CO., AGENTS FOR THE APPLICANTS
Applications Claiming Priority (2)
Application Number | Priority Date | Filing Date | Title |
---|---|---|---|
EPEUROPEANPATENTOFFICE(EPO)10/08/1 | |||
EP89402270 | 1989-08-10 |
Publications (2)
Publication Number | Publication Date |
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IE83457B1 true IE83457B1 (en) | |
IE902911A1 IE902911A1 (en) | 1991-02-27 |
Family
ID=8202984
Family Applications (2)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
IE291190A IE902911A1 (en) | 1989-08-10 | 1990-08-10 | Plants with modified flowers |
IE20020884A IE20020884A1 (en) | 1989-08-10 | 1990-08-10 | Plant with modified flowers |
Family Applications After (1)
Application Number | Title | Priority Date | Filing Date |
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IE20020884A IE20020884A1 (en) | 1989-08-10 | 1990-08-10 | Plant with modified flowers |
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EP (2) | EP0412911B1 (en) |
JP (3) | JP3105242B2 (en) |
AT (2) | ATE203276T1 (en) |
CA (1) | CA2039165C (en) |
DE (2) | DE69034268D1 (en) |
DK (1) | DK0412911T3 (en) |
ES (1) | ES2161681T3 (en) |
FI (1) | FI120156B (en) |
GR (1) | GR3036893T3 (en) |
HK (1) | HK1014738A1 (en) |
HU (1) | HU214927B (en) |
IE (2) | IE902911A1 (en) |
IL (3) | IL95337A (en) |
PT (1) | PT94964B (en) |
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Families Citing this family (228)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
EP0540561B1 (en) * | 1990-07-20 | 1998-11-25 | Pioneer Hi-Bred International, Inc. | Binary cryptocytotoxic method of hybrid seed production |
ATE381622T1 (en) * | 1991-02-07 | 2008-01-15 | Bayer Bioscience Nv | STAMEN SPECIFIC PROMOTORS FROM CORN |
US5770718A (en) * | 1991-05-23 | 1998-06-23 | University Of Waterloo | Gene for APRT from plant tissue |
EP0537399A1 (en) * | 1991-10-16 | 1993-04-21 | Plant Genetic Systems, N.V. | A novel ribonuclease and its inhibitor |
RU2143000C1 (en) * | 1991-11-20 | 1999-12-20 | Моген Интернэшнл Н.В. | Method of preparing plant exhibiting decreased susceptibility to plant parasitic nematodes (variants), recombinant dna (variants), plant transforming vector, strain agrobacterium and method of harvest loss decrease |
EP0577794A4 (en) * | 1992-01-16 | 1994-07-06 | Pacific Seeds Pty Ltd | Genetically modified wheat plants and progeny and method for production of hybrid wheat |
AU676471B2 (en) * | 1992-03-20 | 1997-03-13 | Max-Planck-Gesellschaft Zur Forderung Der Wissenschaften E.V. | Fungus-responsive chimaeric gene |
DE69333880T2 (en) * | 1992-06-12 | 2006-06-22 | Bayer Bioscience N.V. | CONSERVATION OF MALE STERILE PLANTS |
US5750867A (en) * | 1992-06-12 | 1998-05-12 | Plant Genetic Systems, N.V. | Maintenance of male-sterile plants |
EP0665891A1 (en) * | 1992-10-15 | 1995-08-09 | Mogen International N.V. | Genetic moderation or restoration of plant phenotypes |
FR2703561B1 (en) * | 1993-02-15 | 1995-08-04 | Centre Nat Rech Scient | TRANSGENIC PLANTS INCLUDING A TRANSGEN CONSISTING OF A HYBRID NUCLEIC ACID SEQUENCE, COMPRISING AT LEAST ONE FRAGMENT OF UNMITTED MITOCHONDRIAL GENE OF SUPERIOR PLANTS AND PREPARATION. |
GB9401780D0 (en) | 1994-01-31 | 1994-03-23 | Nickerson Biocem Ltd | Modified plants |
US5837850A (en) * | 1994-04-21 | 1998-11-17 | Pioneer Hi-Bred International, Inc. | Regulatory element conferring tapetum specificity |
CA2191441A1 (en) * | 1994-06-06 | 1995-12-21 | Enno Krebbers | Use of anthocyanin genes to maintain male sterile plants |
US7285416B2 (en) | 2000-01-24 | 2007-10-23 | Gendaq Limited | Regulated gene expression in plants |
US7262055B2 (en) | 1998-08-25 | 2007-08-28 | Gendaq Limited | Regulated gene expression in plants |
DE4433975A1 (en) * | 1994-09-23 | 1996-03-28 | Teves Gmbh Alfred | End insulation for electric motor armature |
US5717129A (en) * | 1995-02-16 | 1998-02-10 | Pioneer Hi-Bred International, Inc. | Methods for maintaining sterility in plants |
US6008437A (en) * | 1995-06-06 | 1999-12-28 | Plant Genetic Systems | Use of anthocyanin genes to maintain male sterile plants |
EP0757102A1 (en) | 1995-08-04 | 1997-02-05 | Plant Genetic Systems N.V. | Genetic transformation using a PARP inhibitor |
GB9607517D0 (en) | 1996-04-11 | 1996-06-12 | Gene Shears Pty Ltd | The use of DNA Sequences |
FR2751347B1 (en) * | 1996-07-16 | 2001-12-07 | Rhone Poulenc Agrochimie | CHIMERIC GENE WITH MULTIPLE HERBICIDE TOLERANCE GENES, PLANT CELL AND PLANT TOLERANT WITH MULTIPLE HERBICIDES |
CN1157478C (en) * | 1996-09-03 | 2004-07-14 | 拜尔生物科学股份有限公司 | Improved barster gene |
CA2281862C (en) * | 1997-03-03 | 2014-09-30 | Novartis Ag | Method of hybrid seed production using conditional female sterility |
US6815577B1 (en) | 1997-03-03 | 2004-11-09 | Syngenta Participations Ag | Method of hybrid seed production using conditional female sterility |
US6037523A (en) * | 1997-06-23 | 2000-03-14 | Pioneer Hi-Bred International | Male tissue-preferred regulatory region and method of using same |
CN1206360C (en) | 1998-02-20 | 2005-06-15 | 辛甄塔有限公司 | Hybrid seed production |
KR101085210B1 (en) | 1998-03-20 | 2011-11-21 | 커먼웰쓰 사이언티픽 앤드 인더스트리얼 리서치 오가니제이션 | Control of gene expression |
AUPP249298A0 (en) | 1998-03-20 | 1998-04-23 | Ag-Gene Australia Limited | Synthetic genes and genetic constructs comprising same I |
JP5015373B2 (en) | 1998-04-08 | 2012-08-29 | コモンウェルス サイエンティフィック アンド インダストリアル リサーチ オーガニゼイション | Methods and means for obtaining an improved phenotype |
US8598332B1 (en) | 1998-04-08 | 2013-12-03 | Bayer Cropscience N.V. | Methods and means for obtaining modified phenotypes |
US20040214330A1 (en) | 1999-04-07 | 2004-10-28 | Waterhouse Peter Michael | Methods and means for obtaining modified phenotypes |
US6423885B1 (en) | 1999-08-13 | 2002-07-23 | Commonwealth Scientific And Industrial Research Organization (Csiro) | Methods for obtaining modified phenotypes in plant cells |
US6509516B1 (en) | 1999-10-29 | 2003-01-21 | Plant Genetic Systems N.V. | Male-sterile brassica plants and methods for producing same |
US6506963B1 (en) | 1999-12-08 | 2003-01-14 | Plant Genetic Systems, N.V. | Hybrid winter oilseed rape and methods for producing same |
US6646186B1 (en) * | 2000-07-26 | 2003-11-11 | Stine Seed Farm Inc. | Hybrid soybeans and methods of production |
AU2003215078A1 (en) * | 2002-02-07 | 2003-09-02 | Hybrigene, Inc | Prevention of transgene escape in genetically modified perennials |
US20100031387A1 (en) * | 2002-02-07 | 2010-02-04 | Hybrigene, Inc. | Prevention of transgene escape in genetically modified perennials |
US20030208789A1 (en) * | 2002-05-03 | 2003-11-06 | Stefan Jansens | Wound-inducible expression in plants |
WO2003104464A1 (en) * | 2002-06-06 | 2003-12-18 | Bayer Bioscience N.V. | Female gametophyte specific promoter (zmea1) |
EP1645630A4 (en) * | 2003-06-18 | 2006-09-06 | Japan Tobacco Inc | Method of improving fertility of hybrid plant comprising locating fertility restoration genes on plural loci |
US20060200878A1 (en) | 2004-12-21 | 2006-09-07 | Linda Lutfiyya | Recombinant DNA constructs and methods for controlling gene expression |
CA2620387C (en) | 2005-09-20 | 2018-09-18 | Basf Plant Science Gmbh | Methods for controlling gene expression using ta-sirna |
BRPI0617224A2 (en) | 2005-10-13 | 2011-07-19 | Monsanto Technology Llc | methods for producing hybrid seed |
EP2074227A4 (en) | 2006-10-12 | 2010-03-10 | Monsanto Technology Llc | Plant micrornas and methods of use thereof |
CL2007003743A1 (en) | 2006-12-22 | 2008-07-11 | Bayer Cropscience Ag | COMPOSITION THAT INCLUDES FENAMIDONA AND AN INSECTICIDE COMPOUND; AND METHOD TO CONTROL FITOPATOGENOS CULTURES AND INSECTS FACING OR PREVENTIVELY. |
CL2007003744A1 (en) | 2006-12-22 | 2008-07-11 | Bayer Cropscience Ag | COMPOSITION THAT INCLUDES A 2-PYRIDILMETILBENZAMIDE DERIVATIVE AND AN INSECTICIDE COMPOUND; AND METHOD TO CONTROL FITOPATOGENOS CULTURES AND INSECTS FACING OR PREVENTIVELY. |
US8080688B2 (en) | 2007-03-12 | 2011-12-20 | Bayer Cropscience Ag | 3, 4-disubstituted phenoxyphenylamidines and use thereof as fungicides |
EP1969929A1 (en) | 2007-03-12 | 2008-09-17 | Bayer CropScience AG | Substituted phenylamidines and their use as fungicides |
EP2136627B1 (en) | 2007-03-12 | 2015-05-13 | Bayer Intellectual Property GmbH | Dihalophenoxyphenylamidines and use thereof as fungicides |
EP1969930A1 (en) | 2007-03-12 | 2008-09-17 | Bayer CropScience AG | Phenoxy phenylamidines and their use as fungicides |
EP1969934A1 (en) | 2007-03-12 | 2008-09-17 | Bayer CropScience AG | 4-cycloalkyl or 4-aryl substituted phenoxy phenylamidines and their use as fungicides |
EP2146975B1 (en) | 2007-04-19 | 2015-06-17 | Bayer Intellectual Property GmbH | Thiadiazolyl oxyphenyl amidines and the use thereof as a fungicide |
DE102007045922A1 (en) | 2007-09-26 | 2009-04-02 | Bayer Cropscience Ag | Drug combinations with insecticidal and acaricidal properties |
DE102007045953B4 (en) | 2007-09-26 | 2018-07-05 | Bayer Intellectual Property Gmbh | Drug combinations with insecticidal and acaricidal properties |
DE102007045955A1 (en) | 2007-09-26 | 2009-04-09 | Bayer Cropscience Ag | Active agent combination, useful e.g. for combating animal pests and treating seeds of transgenic plants, comprises substituted amino-furan-2-one compound and at least one compound e.g. diazinon, isoxathion, carbofuran or aldicarb |
DE102007045956A1 (en) | 2007-09-26 | 2009-04-09 | Bayer Cropscience Ag | Combination of active ingredients with insecticidal and acaricidal properties |
DE102007045920B4 (en) | 2007-09-26 | 2018-07-05 | Bayer Intellectual Property Gmbh | Synergistic drug combinations |
DE102007045957A1 (en) | 2007-09-26 | 2009-04-09 | Bayer Cropscience Ag | Active agent combination, useful e.g. for combating animal pests e.g. insects and treating seeds of transgenic plants, comprises substituted amino-furan-2-one compound and at least one compound e.g. benzoyl urea, buprofezin and cyromazine |
DE102007045919B4 (en) | 2007-09-26 | 2018-07-05 | Bayer Intellectual Property Gmbh | Drug combinations with insecticidal and acaricidal properties |
EP2090168A1 (en) | 2008-02-12 | 2009-08-19 | Bayer CropScience AG | Method for improving plant growth |
US8158850B2 (en) * | 2007-12-19 | 2012-04-17 | Monsanto Technology Llc | Method to enhance yield and purity of hybrid crops |
EP2072506A1 (en) | 2007-12-21 | 2009-06-24 | Bayer CropScience AG | Thiazolyloxyphenylamidine or thiadiazolyloxyphenylamidine und its use as fungicide |
EP2168434A1 (en) | 2008-08-02 | 2010-03-31 | Bayer CropScience AG | Use of azols to increase resistance of plants of parts of plants to abiotic stress |
BRPI0918430A2 (en) | 2008-08-14 | 2015-11-24 | Bayer Cropscience Ag | 4-phenyl-1h-pyrazols insecticides. |
DE102008041695A1 (en) | 2008-08-29 | 2010-03-04 | Bayer Cropscience Ag | Methods for improving plant growth |
EP2201838A1 (en) | 2008-12-05 | 2010-06-30 | Bayer CropScience AG | Active ingredient-beneficial organism combinations with insecticide and acaricide properties |
EP2198709A1 (en) | 2008-12-19 | 2010-06-23 | Bayer CropScience AG | Method for treating resistant animal pests |
EP2204094A1 (en) | 2008-12-29 | 2010-07-07 | Bayer CropScience AG | Method for improved utilization of the production potential of transgenic plants Introduction |
EP2039771A2 (en) | 2009-01-06 | 2009-03-25 | Bayer CropScience AG | Method for improved utilization of the production potential of transgenic plants |
EP2039772A2 (en) | 2009-01-06 | 2009-03-25 | Bayer CropScience AG | Method for improved utilization of the production potential of transgenic plants introduction |
EP2039770A2 (en) | 2009-01-06 | 2009-03-25 | Bayer CropScience AG | Method for improved utilization of the production potential of transgenic plants |
EP2387309A2 (en) | 2009-01-19 | 2011-11-23 | Bayer CropScience AG | Cyclic diones and their use as insecticides, acaricides and/or fungicides |
EP2227951A1 (en) | 2009-01-23 | 2010-09-15 | Bayer CropScience AG | Application of enaminocarbonyl compounds for combating viruses transmitted by insects |
JP5592398B2 (en) | 2009-01-28 | 2014-09-17 | バイエル・クロップサイエンス・アーゲー | Disinfectant N-cycloalkyl-N-bicyclic methylene-carboxamide derivatives |
AR075126A1 (en) | 2009-01-29 | 2011-03-09 | Bayer Cropscience Ag | METHOD FOR THE BEST USE OF THE TRANSGENIC PLANTS PRODUCTION POTENTIAL |
EP2218717A1 (en) | 2009-02-17 | 2010-08-18 | Bayer CropScience AG | Fungicidal N-((HET)Arylethyl)thiocarboxamide derivatives |
WO2010094666A2 (en) | 2009-02-17 | 2010-08-26 | Bayer Cropscience Ag | Fungicidal n-(phenylcycloalkyl)carboxamide, n-(benzylcycloalkyl)carboxamide and thiocarboxamide derivatives |
TW201031331A (en) | 2009-02-19 | 2010-09-01 | Bayer Cropscience Ag | Pesticide composition comprising a tetrazolyloxime derivative and a fungicide or an insecticide active substance |
DE102009001469A1 (en) | 2009-03-11 | 2009-09-24 | Bayer Cropscience Ag | Improving utilization of productive potential of transgenic plant by controlling e.g. animal pest, and/or by improving plant health, comprises treating the transgenic plant with active agent composition comprising prothioconazole |
DE102009001681A1 (en) | 2009-03-20 | 2010-09-23 | Bayer Cropscience Ag | Improving utilization of production potential of a transgenic plant by controlling animal pests, phytopathogenic fungi, microorganisms and/or improving plant health, comprises treating plant with a drug composition comprising iprovalicarb |
DE102009001728A1 (en) | 2009-03-23 | 2010-09-30 | Bayer Cropscience Ag | Improving the production potential of transgenic plant, by combating e.g. animal pests and/or microorganism, and/or increasing plant health, comprises treating the plants with active agent composition comprising fluoxastrobin |
DE102009001730A1 (en) | 2009-03-23 | 2010-09-30 | Bayer Cropscience Ag | Improving utilization of production potential of a transgenic plant by controlling animal pests, phytopathogenic fungi and/or microorganisms and/or the plant health, comprises treating plant with a drug composition comprising spiroxamine |
DE102009001732A1 (en) | 2009-03-23 | 2010-09-30 | Bayer Cropscience Ag | Improving the production potential of transgenic plant, by combating e.g. animal pests and/or microorganism, and/or increasing plant health, comprises treating the plants with active agent composition comprising trifloxystrobin |
AU2009342807B2 (en) | 2009-03-25 | 2015-04-02 | Bayer Cropscience Aktiengesellschaft | Synergistic combinations of active ingredients |
CN102448304B (en) | 2009-03-25 | 2015-03-11 | 拜尔农作物科学股份公司 | Active ingredient combinations having insecticidal and acaricidal properties |
EP2232995A1 (en) | 2009-03-25 | 2010-09-29 | Bayer CropScience AG | Method for improved utilisation of the production potential of transgenic plants |
WO2010108508A2 (en) | 2009-03-25 | 2010-09-30 | Bayer Cropscience Ag | Active ingredient combinations with insecticidal and acaricidal properties |
EP2410849A1 (en) | 2009-03-25 | 2012-02-01 | Bayer CropScience AG | Active ingredient combinations having insecticidal and acaricidal properties |
JP5462354B2 (en) | 2009-03-25 | 2014-04-02 | バイエル・クロップサイエンス・アーゲー | Active ingredient combinations with insecticidal and acaricidal properties |
EP2239331A1 (en) | 2009-04-07 | 2010-10-13 | Bayer CropScience AG | Method for improved utilization of the production potential of transgenic plants |
BRPI1015543A8 (en) | 2009-05-06 | 2016-05-24 | Bayer Cropscience Ag | CYCLOPENTANEDIONE COMPOUNDS AND THEIR USE AS INSECTICIDES, ACARICIDES AND/OR FUNGICIDES. |
EP2251331A1 (en) | 2009-05-15 | 2010-11-17 | Bayer CropScience AG | Fungicide pyrazole carboxamides derivatives |
AR076839A1 (en) | 2009-05-15 | 2011-07-13 | Bayer Cropscience Ag | FUNGICIDE DERIVATIVES OF PIRAZOL CARBOXAMIDAS |
EP2255626A1 (en) | 2009-05-27 | 2010-12-01 | Bayer CropScience AG | Use of succinate dehydrogenase inhibitors to increase resistance of plants or parts of plants to abiotic stress |
PL2437595T3 (en) | 2009-06-02 | 2019-05-31 | Bayer Cropscience Ag | Use of fluopyram for controlling sclerotinia ssp |
KR20120051015A (en) | 2009-07-16 | 2012-05-21 | 바이엘 크롭사이언스 아게 | Synergistic active substance combinations containing phenyl triazoles |
WO2011015524A2 (en) | 2009-08-03 | 2011-02-10 | Bayer Cropscience Ag | Fungicide heterocycles derivatives |
EP2292094A1 (en) | 2009-09-02 | 2011-03-09 | Bayer CropScience AG | Active compound combinations |
EP2343280A1 (en) | 2009-12-10 | 2011-07-13 | Bayer CropScience AG | Fungicide quinoline derivatives |
WO2011080256A1 (en) | 2009-12-28 | 2011-07-07 | Bayer Cropscience Ag | Fungicide hydroximoyl-tetrazole derivatives |
US9000012B2 (en) | 2009-12-28 | 2015-04-07 | Bayer Cropscience Ag | Fungicide hydroximoyl-heterocycles derivatives |
EP2519516A2 (en) | 2009-12-28 | 2012-11-07 | Bayer CropScience AG | Fungicidal hydroximoyl-tetrazole derivatives |
BR112012018108A2 (en) | 2010-01-22 | 2015-10-20 | Bayer Ip Gmbh | acaricidal and / or insecticidal combinations of active ingredients |
EP2353387A1 (en) | 2010-02-05 | 2011-08-10 | Bayer CropScience AG | Use of succinate dehydrogenase (SDH) inhibitors in the treatment of plant types in the sweet grass family |
JP2013521255A (en) | 2010-03-04 | 2013-06-10 | バイエル・インテレクチュアル・プロパティ・ゲゼルシャフト・ミット・ベシュレンクテル・ハフツング | Fluoroalkyl-substituted 2-amidobenzimidazoles and their use to enhance stress tolerance in plants |
WO2011113861A2 (en) | 2010-03-18 | 2011-09-22 | Bayer Cropscience Ag | Aryl and hetaryl sulfonamides as active agents against abiotic plant stress |
JP2013523795A (en) | 2010-04-06 | 2013-06-17 | バイエル・インテレクチユアル・プロパテイー・ゲー・エム・ベー・ハー | Use of 4-phenylbutyric acid and / or salt thereof to enhance stress tolerance of plants |
CA2795838A1 (en) | 2010-04-09 | 2011-10-13 | Bayer Intellectual Property Gmbh | Use of derivatives of the(1-cyanocyclopropyl)phenylphosphinic acid, the esters thereof and/or the salts thereof for enhancing the tolerance of plants to abiotic stress |
EP2377397A1 (en) | 2010-04-14 | 2011-10-19 | Bayer CropScience AG | Use of fungicidal agents for controlling mycoses in palm trees |
WO2011134911A2 (en) | 2010-04-28 | 2011-11-03 | Bayer Cropscience Ag | Fungicide hydroximoyl-tetrazole derivatives |
US20130116287A1 (en) | 2010-04-28 | 2013-05-09 | Christian Beier | Fungicide hydroximoyl-heterocycles derivatives |
WO2011134912A1 (en) | 2010-04-28 | 2011-11-03 | Bayer Cropscience Ag | Fungicide hydroximoyl-heterocycles derivatives |
WO2011151370A1 (en) | 2010-06-03 | 2011-12-08 | Bayer Cropscience Ag | N-[(het)arylalkyl)] pyrazole (thio)carboxamides and their heterosubstituted analogues |
EP2576516B1 (en) | 2010-06-03 | 2014-12-17 | Bayer Intellectual Property GmbH | N-[(het)arylethyl)]pyrazole(thio)carboxamides and their heterosubstituted analogues |
UA110703C2 (en) | 2010-06-03 | 2016-02-10 | Байєр Кропсайнс Аг | Fungicidal n-[(trisubstitutedsilyl)methyl]carboxamide |
UA111593C2 (en) | 2010-07-07 | 2016-05-25 | Баєр Інтеллекчуел Проперті Гмбх | ANTRANILIC ACID AMIDES IN COMBINATION WITH FUNGICIDES |
CA2805806A1 (en) | 2010-07-20 | 2012-01-26 | Ruediger Fischer | Use of anthranilic acid amide derivatives for increasing the stress tolerance in plants to abiotic stress |
JP2013532648A (en) | 2010-07-20 | 2013-08-19 | バイエル・クロップサイエンス・アーゲー | Benzocycloalkenes as antibacterial agents |
PL2611300T3 (en) | 2010-09-03 | 2016-10-31 | Substituted annelated dihydropyrimidinone compounds | |
EP2460406A1 (en) | 2010-12-01 | 2012-06-06 | Bayer CropScience AG | Use of fluopyram for controlling nematodes in nematode resistant crops |
BR112013006611B1 (en) | 2010-09-22 | 2021-01-19 | Bayer Intellectual Property Gmbh | method for the control of soy cyst nematode (heterodera glycines) by infesting a nematode resistant soy plant comprising the application of n- {2- [3-chloro-5- (trifluoromethyl) -2-pyridinyl] ethyl} -2 - (trifluoromethyl) benzamide (fluoride |
PE20131399A1 (en) | 2010-10-07 | 2013-12-16 | Bayer Cropscience Ag | FUNGICIDAL COMPOSITION INCLUDING A TETRAZOLILOXIMA DERIVATIVE AND A THIAZOLYLPIPERIDINE DERIVATIVE |
CN103313973B (en) | 2010-10-21 | 2015-09-16 | 拜耳知识产权有限责任公司 | N-benzyl heterocyclic carboxamide |
JP2013541553A (en) | 2010-10-21 | 2013-11-14 | バイエル・インテレクチユアル・プロパテイー・ゲー・エム・ベー・ハー | 1- (Heterocycliccarbonyl) piperidines |
JP2013542215A (en) | 2010-11-02 | 2013-11-21 | バイエル・インテレクチユアル・プロパテイー・ゲー・エム・ベー・ハー | N-hetarylmethylpyrazolyl carboxamides |
EP2640191A1 (en) | 2010-11-15 | 2013-09-25 | Bayer Intellectual Property GmbH | 5-halogenopyrazole(thio)carboxamides |
WO2012065947A1 (en) | 2010-11-15 | 2012-05-24 | Bayer Cropscience Ag | 5-halogenopyrazolecarboxamides |
CN103313971B (en) | 2010-11-15 | 2015-12-02 | 拜耳知识产权有限责任公司 | N-arylpyrazole (sulfo-) methane amide |
EP2460407A1 (en) | 2010-12-01 | 2012-06-06 | Bayer CropScience AG | Agent combinations comprising pyridylethyl benzamides and other agents |
CN103281900A (en) | 2010-12-01 | 2013-09-04 | 拜耳知识产权有限责任公司 | Use of fluopyram for controlling nematodes in crops and for increasing yield |
EP2474542A1 (en) | 2010-12-29 | 2012-07-11 | Bayer CropScience AG | Fungicide hydroximoyl-tetrazole derivatives |
US20130289077A1 (en) | 2010-12-29 | 2013-10-31 | Juergen Benting | Fungicide hydroximoyl-tetrazole derivatives |
EP2471363A1 (en) | 2010-12-30 | 2012-07-04 | Bayer CropScience AG | Use of aryl-, heteroaryl- and benzylsulfonamide carboxylic acids, -carboxylic acid esters, -carboxylic acid amides and -carbonitriles and/or its salts for increasing stress tolerance in plants |
EP2494867A1 (en) | 2011-03-01 | 2012-09-05 | Bayer CropScience AG | Halogen-substituted compounds in combination with fungicides |
CA2823999C (en) | 2011-03-10 | 2020-03-24 | Bayer Intellectual Property Gmbh | Use of lipochito-oligosaccharide compounds for safeguarding seed safety of treated seeds |
CN103502238A (en) | 2011-03-14 | 2014-01-08 | 拜耳知识产权有限责任公司 | Fungicide hydroximoyl-tetrazole derivatives |
BR112013025871A2 (en) | 2011-04-08 | 2016-07-26 | Bayer Ip Gmbh | compound of formula (i) and its use, composition for controlling phytopathogenic fungi, method for controlling phytopathogenic fungi of crops and process for producing compositions |
AR090010A1 (en) | 2011-04-15 | 2014-10-15 | Bayer Cropscience Ag | 5- (CICLOHEX-2-EN-1-IL) -PENTA-2,4-DIENOS AND 5- (CICLOHEX-2-EN-1-IL) -PENT-2-EN-4-INOS REPLACED AS ACTIVE PRINCIPLES AGAINST THE ABIOTIC STRESS OF PLANTS, USES AND TREATMENT METHODS |
EP2511255A1 (en) | 2011-04-15 | 2012-10-17 | Bayer CropScience AG | Substituted prop-2-in-1-ol and prop-2-en-1-ol derivatives |
AR085568A1 (en) | 2011-04-15 | 2013-10-09 | Bayer Cropscience Ag | 5- (BICYCLE [4.1.0] HEPT-3-EN-2-IL) -PENTA-2,4-DIENOS AND 5- (BICYCLE [4.1.0] HEPT-3-EN-2-IL) -PENT- 2-IN-4-INOS REPLACED AS ACTIVE PRINCIPLES AGAINST ABIOTIC STRESS OF PLANTS |
AR085585A1 (en) | 2011-04-15 | 2013-10-09 | Bayer Cropscience Ag | VINIL- AND ALQUINILCICLOHEXANOLES SUBSTITUTED AS ACTIVE PRINCIPLES AGAINST STRIPS ABIOTIQUE OF PLANTS |
HUE043158T2 (en) | 2011-04-22 | 2019-08-28 | Bayer Cropscience Ag | Active compound compositions comprising a (thio)carboxamide derivative and a fungicidal compound |
CN103635483B (en) | 2011-07-01 | 2016-11-09 | 孟山都技术公司 | Method and composition for selective regulation protein expression |
JP2014520776A (en) | 2011-07-04 | 2014-08-25 | バイエル・インテレクチユアル・プロパテイー・ゲー・エム・ベー・ハー | Use of substituted isoquinolinones, isoquinoline diones, isoquinoline triones and dihydroisoquinolinones or their salts in each case as active agents against abiotic stresses in plants |
US9265252B2 (en) | 2011-08-10 | 2016-02-23 | Bayer Intellectual Property Gmbh | Active compound combinations comprising specific tetramic acid derivatives |
CN103748092A (en) | 2011-08-22 | 2014-04-23 | 拜耳知识产权有限责任公司 | Fungicide hydroximoyl-tetrazole derivatives |
EP2561759A1 (en) | 2011-08-26 | 2013-02-27 | Bayer Cropscience AG | Fluoroalkyl-substituted 2-amidobenzimidazoles and their effect on plant growth |
CN103781353B (en) | 2011-09-09 | 2016-10-19 | 拜耳知识产权有限责任公司 | For improveing the acyl homoserine lactones derivant of plant products |
WO2013037717A1 (en) | 2011-09-12 | 2013-03-21 | Bayer Intellectual Property Gmbh | Fungicidal 4-substituted-3-{phenyl[(heterocyclylmethoxy)imino]methyl}-1,2,4-oxadizol-5(4h)-one derivatives |
EA029005B1 (en) | 2011-09-16 | 2018-01-31 | Байер Интеллектчуал Проперти Гмбх | Use of phenylpyrazolin-3-carboxylates for improving plant yield |
AR087872A1 (en) | 2011-09-16 | 2014-04-23 | Bayer Ip Gmbh | USE OF 5-PHENYL-OR 5-BENCIL-2 ISOXAZOLIN-3 CARBOXYLATES TO IMPROVE THE PERFORMANCE OF PLANTS |
AU2012307321B2 (en) | 2011-09-16 | 2016-07-14 | Bayer Intellectual Property Gmbh | Use of acylsulfonamides for improving plant yield |
WO2013041602A1 (en) | 2011-09-23 | 2013-03-28 | Bayer Intellectual Property Gmbh | Use of 4-substituted 1-phenyl-pyrazole-3-carboxylic-acid derivatives as agents against abiotic plant stress |
CN103842507A (en) | 2011-10-04 | 2014-06-04 | 拜耳知识产权有限责任公司 | Rnai for the control of fungi and oomycetes by inhibiting saccharopine dehydrogenase gene |
WO2013050324A1 (en) | 2011-10-06 | 2013-04-11 | Bayer Intellectual Property Gmbh | Combination, containing 4-phenylbutyric acid (4-pba) or a salt thereof (component (a)) and one or more selected additional agronomically active compounds (component(s) (b)), that reduces abiotic plant stress |
MX2014005976A (en) | 2011-11-21 | 2014-08-27 | Bayer Ip Gmbh | Fungicide n-[(trisubstitutedsilyl)methyl]-carboxamide derivatives. |
JPWO2013077420A1 (en) * | 2011-11-25 | 2015-04-27 | 独立行政法人農業生物資源研究所 | Plant transformant, plant transformation method, and vector used in the method |
WO2013079566A2 (en) | 2011-11-30 | 2013-06-06 | Bayer Intellectual Property Gmbh | Fungicidal n-bicycloalkyl and n-tricycloalkyl (thio)carboxamide derivatives |
IN2014CN04325A (en) | 2011-12-19 | 2015-09-04 | Bayer Cropscience Ag | |
CN104039769B (en) | 2011-12-29 | 2016-10-19 | 拜耳知识产权有限责任公司 | 3-[(1,3-thiazole-4-yl methoxyimino) (phenyl) methyl]-2-substituted-1,2,4-diazole-5 (2H) the-one derivant of antifungal |
US9556158B2 (en) | 2011-12-29 | 2017-01-31 | Bayer Intellectual Property Gmbh | Fungicidal 3-[(pyridin-2-ylmethoxyimino)(phenyl)methyl]-2-substituted-1,2,4-oxadiazol-5(2H)-one derivatives |
ES2664230T3 (en) | 2012-02-22 | 2018-04-18 | Bayer Cropscience Ag | Use of fluopiram for the control of diseases of wood in the vine |
CN104244716B (en) | 2012-02-27 | 2017-05-03 | 拜耳知识产权有限责任公司 | Active compound combinations containing a thiazoylisoxazoline and a fungicide |
WO2013139949A1 (en) | 2012-03-23 | 2013-09-26 | Bayer Intellectual Property Gmbh | Compositions comprising a strigolactame compound for enhanced plant growth and yield |
US9357778B2 (en) | 2012-04-12 | 2016-06-07 | Bayer Cropscience Ag | N-acyl-2-(cyclo)alkypyrrolidines and piperidines useful as fungicides |
EP2838893B1 (en) | 2012-04-20 | 2019-03-13 | Bayer Cropscience AG | N-cycloalkyl-n-[(heterocyclylphenyl)methylene]-(thio)carboxamide derivatives |
US20150080337A1 (en) | 2012-04-20 | 2015-03-19 | Bayer Cropscience | N-cycloalkyl-n-[(trisubstitutedsilylphenyl)methylene]-(thio)carboxamide derivatives |
EP2662363A1 (en) | 2012-05-09 | 2013-11-13 | Bayer CropScience AG | 5-Halogenopyrazole biphenylcarboxamides |
EP2662360A1 (en) | 2012-05-09 | 2013-11-13 | Bayer CropScience AG | 5-Halogenopyrazole indanyl carboxamides |
EP2662370A1 (en) | 2012-05-09 | 2013-11-13 | Bayer CropScience AG | 5-Halogenopyrazole benzofuranyl carboxamides |
EP2662362A1 (en) | 2012-05-09 | 2013-11-13 | Bayer CropScience AG | Pyrazole indanyl carboxamides |
JP6262208B2 (en) | 2012-05-09 | 2018-01-17 | バイエル・クロップサイエンス・アクチェンゲゼルシャフト | Pyrazole indanyl carboxamides |
EP2662361A1 (en) | 2012-05-09 | 2013-11-13 | Bayer CropScience AG | Pyrazol indanyl carboxamides |
MX2014013489A (en) | 2012-05-09 | 2015-02-12 | Bayer Cropscience Ag | 5-halogenopyrazole indanyl carboxamides. |
EP2662364A1 (en) | 2012-05-09 | 2013-11-13 | Bayer CropScience AG | Pyrazole tetrahydronaphthyl carboxamides |
AR091104A1 (en) | 2012-05-22 | 2015-01-14 | Bayer Cropscience Ag | COMBINATIONS OF ACTIVE COMPOUNDS THAT INCLUDE A LIPO-CHYTOOLIGOSACARIDE DERIVATIVE AND A NEMATICIDE, INSECTICIDE OR FUNGICIDE COMPOUND |
WO2013184768A1 (en) | 2012-06-05 | 2013-12-12 | University Of Georgia Research Foundation, Inc. | Compositions and methods of gene silencing in plants |
AU2013311826A1 (en) | 2012-09-05 | 2015-03-26 | Bayer Cropscience Ag | Use of substituted 2-amidobenzimidazoles, 2-amidobenzoxazoles and 2-amidobenzothiazoles or salts thereof as active substances against abiotic plant stress |
DK2908641T3 (en) | 2012-10-19 | 2018-04-23 | Bayer Cropscience Ag | PROCEDURE FOR TREATING PLANTS AGAINST FUNGI RESISTANT TO FUNGICIDES USING CARBOXAMIDE OR THIOCARBOXAMIDE DERIVATIVES |
UA114647C2 (en) | 2012-10-19 | 2017-07-10 | Байєр Кропсайнс Аг | Method for enhancing tolerance to abiotic stress in plants using carboxamide or thiocarboxamide derivatives |
US9801374B2 (en) | 2012-10-19 | 2017-10-31 | Bayer Cropscience Ag | Active compound combinations comprising carboxamide derivatives |
JP6262747B2 (en) | 2012-10-19 | 2018-01-17 | バイエル・クロップサイエンス・アクチェンゲゼルシャフト | Plant growth promotion method using carboxamide derivatives |
EP2735231A1 (en) | 2012-11-23 | 2014-05-28 | Bayer CropScience AG | Active compound combinations |
UA116222C2 (en) | 2012-11-30 | 2018-02-26 | Байєр Кропсайєнс Акцієнгезелльшафт | Ternary fungicidal and pesticidal mixtures |
UA116223C2 (en) | 2012-11-30 | 2018-02-26 | Байєр Кропсайєнс Акцієнгезелльшафт | Binary fungicidal mixtures |
BR112015012055B1 (en) | 2012-11-30 | 2021-01-12 | Bayer Cropscience Ag | ternary fungicidal composition, its preparation process, method to control one or more harmful microorganisms, seed resistant to harmful microorganisms and its treatment method |
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TW201507722A (en) | 2013-04-30 | 2015-03-01 | Bayer Cropscience Ag | N-(2-halogen-2-phenethyl)carboxamides as nematicides and endoparasiticides |
WO2014177514A1 (en) | 2013-04-30 | 2014-11-06 | Bayer Cropscience Ag | Nematicidal n-substituted phenethylcarboxamides |
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US20210323950A1 (en) | 2018-06-04 | 2021-10-21 | Bayer Aktiengesellschaft | Herbicidally active bicyclic benzoylpyrazoles |
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JP2022500459A (en) | 2018-09-17 | 2022-01-04 | バイエル・アクチエンゲゼルシヤフト | Use of the fungicide isofukusiplum for the control of ergot in grains and the reduction of sclerotia |
WO2020058144A1 (en) | 2018-09-17 | 2020-03-26 | Bayer Aktiengesellschaft | Use of the succinate dehydrogenase inhibitor fluopyram for controlling claviceps purpurea and reducing sclerotia in cereals |
Family Cites Families (12)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
US4956282A (en) * | 1985-07-29 | 1990-09-11 | Calgene, Inc. | Mammalian peptide expression in plant cells |
FI875467A (en) * | 1986-12-19 | 1988-06-20 | Agricultural Genetics Co | DNA MOLEKYLER, SOM AER NYTTIGA VID VAEXTSKYDD. |
GB8901677D0 (en) * | 1989-01-26 | 1989-03-15 | Ici Plc | Hybrid seed production |
NZ224787A (en) * | 1987-05-26 | 1990-08-28 | Calgene Inc | A dna construct for specifically modifying the phenotype of fruit as distinct from other plant tissue |
US5356799A (en) * | 1988-02-03 | 1994-10-18 | Pioneer Hi-Bred International, Inc. | Antisense gene systems of pollination control for hybrid seed production |
NZ227835A (en) * | 1988-02-03 | 1992-09-25 | Paladin Hybrids Inc | Antisense gene systems of pollination control for hybrid seed production |
DE3810286A1 (en) * | 1988-03-25 | 1989-10-12 | Max Planck Gesellschaft | TRANSGENIC PLANT WITH MODIFIED PHYSIOLOGY, MORPHOLOGY AND MODIFIED HORMONE METABOLISM, TISSUE CULTURES OF THIS PLANT AND METHOD FOR THE PRODUCTION THEREOF |
GB8810120D0 (en) * | 1988-04-28 | 1988-06-02 | Plant Genetic Systems Nv | Transgenic nuclear male sterile plants |
ATE195218T1 (en) * | 1988-06-20 | 2000-08-15 | Novartis Erfind Verwalt Gmbh | METHOD FOR CONTROLLING PLANT PESTS USING NON-VEGETABLE PROTEINASE INHIBITORS |
DE69034190T2 (en) * | 1989-02-02 | 2006-02-23 | Pioneer Hi-Bred International, Inc. | MOLECULAR PROCESSES FOR THE MULTIPLICATION OF HYBRID SEEDS |
EP0412676B1 (en) * | 1989-08-07 | 1995-06-07 | Yissum Research Development Company Of The Hebrew University Of Jerusalem | Method for producing the spiroplasma SP. DNA methylase |
WO1991009957A1 (en) * | 1989-12-22 | 1991-07-11 | E.I. Du Pont De Nemours And Company | Site-specific recombination of dna in plant cells |
-
1990
- 1990-08-09 DE DE69034268T patent/DE69034268D1/en not_active Expired - Lifetime
- 1990-08-09 DK DK90402281T patent/DK0412911T3/en active
- 1990-08-09 EP EP90402281A patent/EP0412911B1/en not_active Expired - Lifetime
- 1990-08-09 WO PCT/EP1990/001315 patent/WO1991002069A1/en active Application Filing
- 1990-08-09 JP JP02510866A patent/JP3105242B2/en not_active Expired - Lifetime
- 1990-08-09 AT AT90402281T patent/ATE203276T1/en not_active IP Right Cessation
- 1990-08-09 DE DE69033764T patent/DE69033764T2/en not_active Expired - Lifetime
- 1990-08-09 ES ES90402281T patent/ES2161681T3/en not_active Expired - Lifetime
- 1990-08-09 AT AT00128293T patent/ATE496135T1/en not_active IP Right Cessation
- 1990-08-09 HU HU906172A patent/HU214927B/en unknown
- 1990-08-09 CA CA002039165A patent/CA2039165C/en not_active Expired - Lifetime
- 1990-08-09 EP EP00128293A patent/EP1090999B1/en not_active Expired - Lifetime
- 1990-08-10 IE IE291190A patent/IE902911A1/en not_active IP Right Cessation
- 1990-08-10 IL IL95337A patent/IL95337A/en active IP Right Grant
- 1990-08-10 ZA ZA906344A patent/ZA906344B/en unknown
- 1990-08-10 IE IE20020884A patent/IE20020884A1/en not_active IP Right Cessation
- 1990-08-10 IL IL12130190A patent/IL121301A/en active IP Right Grant
- 1990-08-10 PT PT94964A patent/PT94964B/en not_active IP Right Cessation
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1991
- 1991-04-10 FI FI911728A patent/FI120156B/en active IP Right Grant
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1996
- 1996-12-04 JP JP32389896A patent/JP3720497B2/en not_active Expired - Lifetime
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1997
- 1997-07-14 IL IL12130197A patent/IL121301A0/en unknown
-
1998
- 1998-02-10 US US09/021,785 patent/US6046382A/en not_active Expired - Fee Related
- 1998-12-28 HK HK98116145A patent/HK1014738A1/en not_active IP Right Cessation
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1999
- 1999-07-21 JP JP20686499A patent/JP3609654B2/en not_active Expired - Lifetime
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2001
- 2001-10-16 GR GR20010401763T patent/GR3036893T3/en not_active IP Right Cessation
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