WO2012065219A1 - Drought tolerant plants - Google Patents
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- WO2012065219A1 WO2012065219A1 PCT/AU2011/001478 AU2011001478W WO2012065219A1 WO 2012065219 A1 WO2012065219 A1 WO 2012065219A1 AU 2011001478 W AU2011001478 W AU 2011001478W WO 2012065219 A1 WO2012065219 A1 WO 2012065219A1
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
- A01H1/12—Processes for modifying agronomic input traits, e.g. crop yield
- A01H1/122—Processes for modifying agronomic input traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- A01H1/1225—Processes for modifying agronomic input traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for drought, cold or salt resistance
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
- A01H1/04—Processes of selection involving genotypic or phenotypic markers; Methods of using phenotypic markers for selection
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H5/00—Angiosperms, i.e. flowering plants, characterised by their plant parts; Angiosperms characterised otherwise than by their botanic taxonomy
- A01H5/10—Seeds
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- C07—ORGANIC CHEMISTRY
- C07K—PEPTIDES
- C07K14/00—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof
- C07K14/415—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from plants
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- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
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- 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
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- 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
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- 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
- C12N15/8262—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield involving plant development
- C12N15/8266—Abscission; Dehiscence; Senescence
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- 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
- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- C12N15/8273—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for drought, cold, salt resistance
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- Y—GENERAL TAGGING OF NEW TECHNOLOGICAL DEVELOPMENTS; GENERAL TAGGING OF CROSS-SECTIONAL TECHNOLOGIES SPANNING OVER SEVERAL SECTIONS OF THE IPC; TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
- Y02—TECHNOLOGIES OR APPLICATIONS FOR MITIGATION OR ADAPTATION AGAINST CLIMATE CHANGE
- Y02A—TECHNOLOGIES FOR ADAPTATION TO CLIMATE CHANGE
- Y02A40/00—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production
- Y02A40/10—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production in agriculture
- Y02A40/146—Genetically Modified [GMO] plants, e.g. transgenic plants
Definitions
- FIELD FIELD
- the present specification teaches the generation of drought tolerant plants.
- the present disclosure enables manipulation of a phenotypic characteristic referred to herein as "stay-green" to generate drought tolerant plants by recombinant, mutagenic and/or breeding and selection methods.
- Plant management practice systems to increase crop yield and harvest efficiency in water-limited environments are also taught herein.
- Drought is the single most important constraint to cereal production worldwide. Sorghum is a repository of drought resistance mechanisms, which include C 4 photosynthesis, deep roots and thick leaf wax which enable growth in hot and dry environments.
- Drought tolerance makes sorghum especially important in dry regions such as sub-Saharan Africa, western-central India, north-eastern Australia, and the southern plains of the US. With increasing pressure on the availability of scarce water resources, the identification of traits associated with grain yield under drought conditions becomes more important.
- grain yield is a function of transpiration (T), transpiration efficiency (TE), and harvest index (HI) [Passioura, J. Aust. Inst. Agric. Sci. 43:117-120, 1977].
- T transpiration
- TE transpiration efficiency
- HI harvest index
- grain yield is linked to post-anthesis T (Turner, J. Exp. Bot. 55:2413-2425, 2004; Manschadi et al, Fund Plant. Biol 53:823-837, 2006), because HI increases with the fraction of total crop T used after anthesis (Passioura, 1977 supra; Sadras and Connor, Field Crops Res. 26:227-239, 1991 ; Hammer, Agric. Sci. J 9:16-22, 2006).
- post-anthesis T is associated with reduced drought stress around anthesis, which can positively affect crop growth rate at anthesis of cereals and hence grain number (Andrade et al, Crop Sci. 42:1173-1179, 2002; Van Oosterom and Hammer, Field Crops Res. 108:259-268, 2008).
- post-anthesis T can be increased by restricting pre-anthesis T. This can be achieved by restricting canopy size, either genetically or through crop management. However, a smaller canopy will only reduce total T if its TE is not compromised.
- Significant genotypic differences in TE have been reported for sorghum (Hammer et al, Aust. J. Agric. Res.
- post-anthesis water use can be increased by increasing the total amount of water accessed by the crop, either through deeper rooting or reduced lower limit of water extraction (Manschadi et al, 2006 supra).
- the stay-green trait affects a number of the above processes in sorghum. First, stay- green reduces water use during the pre-anthesis period by restricting canopy size (via reduced tillering and smaller leaves).
- stay-green improves water accessibility by increasing the root: shoot ratio. There is some experimental evidence for better water extraction in stay-green lines, although more research is required. These root responses could also be explained by enhanced auxin transport (Wang et al, Molecular Plant 2(3 ⁇ 4):823-831, 2009).
- stay- green increases the greenness of leaves at anthesis, effectively increasing photosynthetic capacity, and, therefore, TE (providing that photosynthesis increases proportionately more than conductance). The increase in leaf greenness is an indirect affect of reduced leaf mass, i.e. nitrogen is concentrated in the leaf.
- the present disclosure teaches quantitative trait loci (QTLs) associated with and/or which otherwise facilitate the stay-green phenotype.
- QTLs quantitative trait loci
- the QTLs are referred to herein as "stay-green (Stg) X" wherein X is a numeral increasing from 1 which represents the region • on a chromosome associated with the stay-green phenotype.
- the QTLs identify genetic regions on sorghum which carry loci ' which encode one to a number of proteins or regulatory agents such as microRNAs which facilitate the stay-green phenotype. Modulation of expression of one or more these loci in a crop plant generates a canopy architecture which facilitates a shift in water use by the plant to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency thereby increasing harvest index (HI) and grain yield . • under water-limited conditions.
- the loci encode a PIN protein which is associated with an auxin.
- PIN proteins are auxin efflux carriers which contain transmembrane domains and are mainly localized in the plasma membranes.
- the term "PIN” is derived from "pin-like inflorescence".
- SbPINn is used to describe such a gene in sorghum wherein n is a numeral defining the auxin efflux carrier component and n is 1 through 1 1.
- Reference to “SbPINn” includes its homologs and orthologs in other plants. Examples of SbPINn loci in sorghum include those listed in Table 1 A such as but not limited to SbPIN4 and SbPIN2 or their equivalents in other plants. Modulation of expression of a ⁇ or expression of a PIN with a particular polymorphic variation is taught herein to facilitate exhibition of the stay-green phenotype.
- the present disclosure teaches PINs from other plants such as rice.
- the letter prefix has a PIN specifies its source (eg Sb, sorghum; Os, rice; etc).
- the location of a SbPIN in sorghum is determined by gene ID number (See Table 1A).
- SbPIN4 corresponds to OsPIN5
- SbPIN2 corresponds to OsPIN3a.
- SbPIN4 and SbPIN2 are examples of SbPINs taught herein to be responsible for the stay-green trait in sorghum associated with Stgl, and Stg2, respectively, resulting in a range of phenotypes that confer drought adaptation via decreased water use until anthesis (due to reduced tillering and smaller leaves), increased water accessibility (due to enhanced roo shoot ratio), increased transpiration efficiency under mild water deficit (due to higher leaf nitrogen concentration), increased biomass per leaf area under terminal water deficit (due to increased transpiration per leaf area) and increased grain yield, grain size and lodging resistance.
- Other examples are listed in Table 1A and include their equivalents in other plants.
- the present disclosure teaches a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising modulating the level of expression of an existing or introduced pin-like inflorescence (PIN) locus in all or selected tissue in the plant to facilitate a stay-green phenotype, which phenotype includes a shift in water use to the post- anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water- limited conditions.
- PIN pin-like inflorescence
- a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant or parent of the plant a genetic agent encoding a PIN protein selected from SbPINl to 11 or an equivalent thereof from another plant or functional homolog or ortholog thereof; or which modulates expression of an indigenous PIN protein; wherein the level and location of PIN expression facilitate a stay-green phenotype, which phenotype includes, inter alia, a canopy architecture which facilitates a shift in water use to the post-anthesis period or increased accessibility of water during crop growth resulting in increased harvest index and grain yield under water limiting conditions.
- the present disclosure further teaches a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant or parent of the plant a genetic agent which encodes a sorghum PIN protein selected from SbPIN4 and SbPIN2 or an equivalent in another plant or which modulate expression of an indigenous PIN.
- a genetic agent which encodes a sorghum PIN protein selected from SbPIN4 and SbPIN2 or an equivalent in another plant or which modulate expression of an indigenous PIN.
- Reference to modulating includes increasing and decreasing the level of expression.
- a PIN may be selected having a desired expression profile in all or selected tissues in a plant.
- a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species comprising introducing into the plant or a parent of the plant a genetic agent which encodes a product which is associated with or facilitates a stay-green phenotype which phenotype includes a shift in water use to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water-limited conditions, and wherein the product is selected from the list consisting of SbPINl to 11 or an equivalent thereof in another plant or which agent modulates expression of an indigenous PIN to the plant.
- Also taught herein is a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant or parent of the plant a genetic agent which encodes a protein selected from the list consisting of SbPIN4 and SbPIN2 or an equivalent thereof in another plant or which agent modulates expression of an indigenous PIN in the plant.
- introducing includes via recombinant intervention as well as by mutagenesis or breeding protocol followed by selection.
- the instant disclosure is instructional for a method of generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant and/or parent of the plant a genetic agent which encodes two or more PIN proteins or which modulates expression of two or more indigenous PIN proteins in a plant.
- PINs in sorghum include SbPINl to 1 1 such as SbPIN4 and SbPIN2 or an equivalent thereof from another plant.
- SbPIN4 corresponds to OsPIN5
- SbPIN2 corresponds to OsPIN3a.
- SbPINs and OsPINS are defined in Table 1 A.
- Genetic material which encodes a PIN protein or functional homolog or ortholog thereof which is associated with or facilitates a stay-green phenotype, which phenotype includes a canopy architecture which facilitates a shift in water use to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water limiting conditions and an agent which modulates expression of the PIN; are both enabled herein.
- the genetic material is selected from (i) an agent which encodes SbPIN4; and ( ⁇ ) an agent which modulates levels of expression of SbPIN4 or its equivalent in another plant.
- the genetic material selected from (i) an agent which encodes SbPIN2; and (ii) an agent which modulates expression of SbPIN2 or its equivalent in another plant.
- the genetic material includes an agent which encodes a SbPIN4 or SbPIN2 protein or functional homolog or ortholog thereof or an equivalent thereof in another plant which is associated with or facilitates a stay-green phenotype, which phenotype includes a canopy architecture which facilitates a shift in water use to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water limiting conditions; and an agent which modulates expression of SbPIN4 or SbPIN2, or an equivalent thereof in another plant
- the plant management system includes the generation of drought adapted crop including cereal plants using the selection and modulation of expression of a PIN locus or its functional equivalent as herein defined alone or in combination with the introduction of other useful traits such as grain size, root size, salt tolerance, herbicide resistance, pest resistance and the like.
- the plant management system comprises generation of drought adapted plants and agricultural procedures such as irrigation, nutrient requirements, crop density and geometry, weed control, insect control, soil aeration, reduced tillage, raised beds and the like.
- Examples of a PIN locus include SbPIN 1 to 11 (Table 1A) such as SbPIN4 and SbPIN2 and an equivalents thereof in another plant.
- a business model is also taught herein for improved economic returns on crop yield, the model comprising generating crop plants having a canopy architecture which facilitates a shift in water use by the plant to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency thereby increasing HI and grain yield under water-limited conditions, obtaining seed from the generated crop plant and' distributing the seed to grain producers to enhance yield and profit.
- HW high water HWHD
- high water high density (intermediate water stress)
- HWLD high water, low density (least water stressed)
- PASM post-anthesis stem mass
- Figure 1 is a graphical representation showing the relation between culms per m 2 and green leaf area at anthesis in a range of near-isogenic lines containing various Stg introgressions.
- Figure 3 is a graphical representation showing the histogram of predicted values for culms per plant in the Stgl fine-mapping population averaged over three seasons.
- Figure 4 is a tabulated representation showing a histogram of culms per plant at 44 DAE for five genotypes grown under two water regimes.
- the genotypes comprise RTx7000 (recurrent parent), 6078-1 (donor parent), and three selections from the Stgl fine-mapping population.
- HWLD high water, low density (10 plants/m 2 ).
- LWLD low water, low density (10 plants/m 2 ).
- Figure 5 is a graphical representation showing the phenotypic variation in the Stgl fine-mapping population for presence of T2.
- Figure 6 is a graphical representation showing the phenotypic variation in the Stgl fine-mapping population for presence of T3.
- Figure 7 is a representation showing a histogram of T2 presence for eight high- tillering and eight low-tillering recombinants from the Stgl fine-mapping population.
- Figure 8 is a representation showing a histogram of total tiller number per plant for five high-tillering and three low-tillering recombinants from the Stgl fine-mapping population. A value of 2.5 was chosen as the arbitrary cut-off between high and low tillering.
- Figures 9A through D are graphical representations showing the leaf size distribution of mainstem and tillers for RTx7000 and 6078-1 (Stgl NIL) grown in lysimeters under low and high VPD conditions.
- Figure 10 is a graphical representation showing the mainstem leaf size
- Figure 11 is a graphical representation showing the leaf size distribution (Ll-6) for the parents of the Stgl fine-mapping population grown in an igloo.
- Figure 12 is a graphical representation showing the leaf length distribution (Ll-6) for the parents of the Stgl fine-mapping population grown in an igloo.
- Figure 13 is a graphical representation showing the leaf width distribution (Ll-6) for the parents of the Stgl fine-mapping population grown in an igloo.
- Figure 14 is a graphical representation showing the leaf size distribution (11-11) for the parents of the Stgl fine-mapping population grown in an igloo.
- Figure 15 is a graphical representation showing the leaf length distribution (LI -10) for the parents of the Stgl fine-mapping population grown in an igloo.
- Figure 16 is a representation showing a histogram of phenotypic variation for L10 length in a subset of the Stgl fine-mapping population grown in an igloo.
- Figure 17 is a diagrammatic representation showing that increased water availability at anthesis is achieved via reduced water use due to two mechanisms (reduced tillering and smaller leaves) in plants containing the Stgl region.
- Figure 18 is a representation showing that canopy size is modulated by both constitutive and adaptive responses controlled by a gene(s) in the Stgl region.
- Figure 20 is a graphical representation showing the relation between the area of leaf 12 and the total green leaf area at anthesis for the two parents (6078-1 and RTx7000) and three recombinants from the Stgl fine-mapping population.
- Figure 21 is a graphical representation showing the relation between total green leaf area (cm 2 /m 2 ) and crop water use (mm) at anthesis for the two parents (6078-1 and RTx7000) and three recombinants from the Stgl fine-mapping population.
- Figure 22 is a graphical representation showing the relation between green leaf area and water use (T) in four Stg QTL and the recurrent parent (RTx7000) in lysimetry studies under two levels of VPD.
- Figure 23 is a graphical representation showing a histogram of phenotypic variation for the "roo shoot ratio" at L6 in the Stgl fine-mapping population grown in an igloo.
- Figure 24 is a graphical representation showing the temporal pattern of cumulative crop water use for RTx7000 and Stgl grown under low- water and low-density (20 plants/m 2 ) conditions. The vertical line marks anthesis.
- Figure 25 is a graphical representation showing the relation between the length (mm) and greenness (SPAD) of leaf 10 in the Stgl fine-mapping propulation grown in an igloo.
- Figure 26 is a graphical representation showing the relation between leaf greenness (SPAD) and leaf photosynthesis in a subset of lines from the Stgl fine-mapping population, including the parents.
- SPAD leaf greenness
- Figure 27 is a graphical representation showing the relation between leaf greenness (SPAD) and WUE (Licor) in a subset of lines from the Stgl fine-mapping population, including the parents.
- Figure 28 is a graphical representation showing the relation between leaf greenness (SPAD) and WUE (Licor) in four Stg Nils (Stgl , Stg2, Stg3 and Stg4) and the recurrent parent (RTx7000).
- Figure 29 is a graphical representation showing the relation between transpiration per leaf area and transpiration efficiency in four Stg QTL and the recurrent parent (RTx7000) in lysimetry studies under two levels of VPD.
- Figure 30 is a graphical representation showing the relation between CWU (mm) before and after anthesis in a subset of lines from the Stgl fine-mapping population, including the parents, grown under high density (HD) and low density (HD) conditions.
- Figures 31 A and B are graphical representations showing patterns of cumulative water use for Stgl and RTx7000 grown under LWHD and conditions.
- Figure 32 is a graphical representation showing the relation between CWU (mm) before and after anthesis in four Stg QTL and the recurrent parent (RTx7000) grown under low water (LW) and low density (LD) conditions.
- PASM post-anthesis stem mass
- RTx7000 recurrent parent
- RWC relative water content
- FL-2 mid-grain filling
- RTx7000 recurrent parent
- Figures 56A through C are graphical representations showing results from running a sorghum crop simulation model using the generic variety Buster with the usual 2 tillers/plant (HT) versus a Buster with only 1 tiller/plant (LT) in a well-watered (WW) and a terminally stressed (TS) virtual environment.
- Figure 57A is a graphical representation of differential expression of SbPIN4 (Stgl candidate) under well-watered conditions. Under well-watered conditions, this gene is down-regulated in young root tips Tx642 and Stgl NIL compared to Tx7000.
- Figure 57B is a graphical representation of differential expression of SbPIN4 (Stgl candidate) under water-deficient conditions.
- FIG. 57C is a graphical representation of differential expression of SbPTN2 (Stg2 candidate) under well-watered conditions. Under well-watered conditions, this gene is slightly up-regulated in stem and root tissues of Tx642 and Stgl NIL compared to Tx7000.
- Figure 57D is a graphical representation of a differential expression of SbPlN2 (Stg 2 candidate) under water-deficient conditions. Under water-deficient conditions, this gene is up-regulated in most tissues of Tx642 and Stgl NIL compared to Tx7000.
- the present disclosure teaches QTLs associated with and which facilitate the stay- green phenotype in crop including cereal plants.
- the QTLs are referred generically as StgX wherein X is a numeral from 1 and above corresponding to a genetic locus or genetic loci region on a particular chromosome in a crop plant.
- a sub-region is referred to as StgXm where m is an alphabetical designation of a region within StgX.
- Modulation of expression of an StgX in all or selected tissue in a plant is taught herein to facilitate a physiological and genetic network which induces or promotes a shift in water use by the crop plant to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency, thereby increasing harvest index (HI) and grain yield under water-limited conditions.
- "Expression" of an StgX includes up- regulating or down-regulating expression levels (i.e. modulation of expression) of a locus as well as selection of a polymorphic variant which is expressed at a higher or lower level or which encodes a more or less active product in all or selected tissue in a plant.
- the locus may itself confer this phenotype or a functional equivalent thereof such as a cDNA encoding the same protein encoded by the locus.
- the QTL's identify loci encoding PIN proteins.
- PIN proteins are auxin efflux carriers which contain transmembrane domains and are mainly localized in the plasma membranes.
- the term "PIN” is derived from "pin-like inflorescence".
- a pin for sorghum is "SbPIN”.
- the instant disclosure teaches a PIN from any plant (e.g OsPIN from rice). The genomic location of an SbPIN from sorghum is described in Table 1 A.
- a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant or parent of the plant a genetic agent encoding a PIN protein or functional homolog or ortholog thereof; or which modulates expression of an indigenous PIN protein; wherein the level and location of PIN expression facilitate a stay-green phenotype, which phenotype includes, inter alia, a canopy architecture which facilitates a shift in water use to the post-anthesis period or increased accessibility of water during crop growth resulting in increased harvest index and grain yield under water limiting conditions.
- the present disclosure further teaches a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant or parent of the plant a genetic agent which encodes a sorghum SbPIN protein selected from SbPIN 1 through SbPIN 11 or an equivalent in another plant or which modulate expression of an indigenous PIN.
- PINs include SbPIN4 and SbPIN2 and other SbPINs listed in Table 1A and their equivalents in another plant as well as a PIN having a particularly desired polymorphic variation which, for example, permits an altered expression profile of elevated levels of the PIN protein.
- the genetic agent may be a locus or genomic region or its functional equivalent such as cDNA or genomic DNA fragment. Alternatively the agent may modulate expression as an indigenous PIN locus in a particular plant.
- introducing means by recombinant intervention, or by mutagenisis or breeding followed by selection.
- modulation expression of a PIN alone or in combination with a genetic or physiological network alters plant architecture to enhance or otherwise promote efficient water use.
- the modified architecture is modified plant canopy architecture.
- progeny includes immediate progeny as well as distant relatives of the plant, as long as it stably expresses the stay-green trait first introduced to an earlier parent.
- Reference to a "crop plant” includes a cereal plant.
- the crop plants enabled herein include sorghum, wheat, oats, maize, barley, rye, rice, abaca, alfalfa, almond, apple, asparagus, banana, bean-phaseolus, blackberry, broad bean, canola, cashew, cassava, chick pea, citrus, coconut, coffee, corn, cotton, fig, flax, grapes, groundnut, hemp, kenaf, lavender, mano, mushroom, olive, onion, pea, peanut, pear, pearl millet, potato, ramie, rapseed, ryegrass, soybean, strawberry, sugarbeet, sugarcane, sunflower, sweetpotato, taro, tea, tobacco, tomato, triticale, truffle and yam.
- the drought tolerance mechanisms of sorghum are used to promote drought tolerance in sorghum as well as other crop plants.
- the genetically modified plant uses water more efficiently than a non-genetically modified plant of the same species.
- Existing PIN loci in each of the above plants are referred to as "indigenous" PINs.
- the instant disclosure teaches up- and down-regulating an indigenous PIN or selecting a PIN having a particular expression profile.
- An "indigenous" PIN is a PIN locus in a parent plant prior to any manipulation (recombinant, mutagenisis or breeding).
- drought tolerance includes drought escape, drought adaptation, drought resistance, reduced sensitivity to drought conditions, enhanced water use efficiency as well as an ability to shift water use to the post-anthesis period or increased accessibility of water during crop growth, thereby increasing HI and grain yield under water-limited conditions.
- Plants exhibiting drought tolerance are described as “drought adapted plants” or “plants exhibiting reduced sensitivity to water-limited conditions”. Taught herein is that drought tolerance is induced, facilitated by or otherwise associated with the stay-green phenotype.
- genetically modified in relation to a plant, includes an originally derived genetically modified plant as well as any progeny, immediate or distant which stably express the stay-green trait.
- the present disclosure teaches both classical breeding techniques to introduce the genetic agent, i.e. the stay-green QTL or a functional equivalent thereof such as cDNA or a genomic fragment or an agent which up-regulates or down-regulates (i.e. modulates) expression of the QTL or the protein encoded therefrom as well as genetic engineering technology.
- the latter is encompassed by the terms “genetic engineering means” and "recombinant means”.
- Markers defining stay-green can also be screened during breeding protocols to monitor transfer of particular genetic regions.
- a specific stay-green region is genetically inserted by recombinant means into a plant cell or plant callus and a plantlet regenerated.
- a "genetically modified" plant includes a parent or any progeny as well as any products of the plant such as grain, seed, propagating material, pollen and ova.
- a PIN locus may be expressed in one particular plant tissue but not expressed or its expression reduced in another tissue.
- a plant may be subject to mutagenisis such as genetic, radioactive or chemical mutagenisis and mutated plants selected with a PIN having a desired phenotype.
- Reference to the "stay-green phenotype" includes characteristics selected from enhanced canopy architecture plasticity, reduced canopy size, enhanced biomass per unit leaf area at anthesis, higher transpiration efficiency, increased water use during grain filling, increased plant water status during grain filling, reduced pre:post anthesis biomass ratio, delayed senescence, increased grain yield, larger grain size, and reduced lodging.
- a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species comprising introducing into the plant or a parent of the plant a genetic agent which encodes a product which is associated with or facilitates a stay-green phenotype which phenotype includes a shift in water use to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water-limited conditions, and wherein the product is selected from the list consisting of SbPINl to 11 including SbPIN4 and SbPIN2, and other SbPINs listed in Table 1 A or an equivalent thereof in another plant or which agent modulates expression of an indigenous PIN in the plant.
- Enabled herein ⁇ are genetically modified plants which exhibit the stay-green phenotype as a result of the genetic modification as well as seeds, fruit, flowers and other reproductive or other propagating material. Also enabled are root stock and propagating stock. This is based on the premise that the seeds, fruit, flowers, reproductive and propagating material exhibit or can pass on the stay-green phenotype introduced into the ultimate parent(s).
- Reference to an "agent which modulates levels of expression of a PIN” includes promoters, microRNAs, genes and chemical compounds which facilitate increased or decreased expression of the gene in all or selected tissue or increased or decreased activity of a gene product as well as cDNA and genomic fragments.
- An agent may also be an intron of a genomic gene which is part of a natural genetic network to facilitate modulation of expression.
- a PIN protein produces an auxin gradient in cells and contains transmembrane domain and is mainly localized in the plasma membrane.
- PIN proteins are the rate limiting ' factors of auxin transport and provide vectorial direction for the auxin flows. It is taught herein that at least one of Stgl or Stg2 encodes a PIN protein. Introduction of Stgl or Stg2 de novo in a plant or elevation of its modulation or expression of an indigenous Stgl or Stg2 is taught to facilitate exhibition of one or more features or sub-features associated with the stay-green phenotype.
- PIN proteins are efflux carriers of auxin which mediate polar auxin transport (PAT) from cell to cell as opposed to the transport of auxin through the xylem (Rashotte et al.. Plant Cell 13:1683-1697, 2000; Friml et al, Current Opinion in Plant Biology 6:7-12, 2003).
- the term 'PIN' is derived from the pin-like inflorescence which develop in Arabidopsis when auxin transport is defective.
- SbPINn from sorghum where n is a numeral from 1 through 1 1 as well as a PIN from any other plant.
- Also taught herein is a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant or parent of the plant a genetic agent which encodes a protein selected from the list consisting of SbPINl to 1 1 such as SbPIN4 and SbPIN2 or other SbPINs listed in Table 1 A or an equivalent thereof in another plant or which agent modulates expression of an indigenous PIN in the plant.
- sorghum SbPIN4 and SbPIN2 are major drought adaptation genes along with other SbPINs as well as their equivalents in other plants. Differences in auxin signalling explain all of the multiple phenotypes observed in plants with a PIN expression profile. Phenotypes exhibited by SbPIN4 or SbPIN2 plants for example are explained by changes in auxin efflux and include reduced tillering, smaller leaves (both length and width), reduced leaf mass and increased roo shoot ratio.
- Phenotypes exhibited by SbPIN4 or SbPIN2 plants for example can also be explained indirectly (or as emergent consequences of these direct effects) and include increased availability of water at anthesis, higher leaf N concentration at anthesis, increased transpiration and biomass per unit leaf area, higher transpiration efficiency, retention of green leaf area during grain filling, increased harvest index, higher grain yield, larger grain size and increased lodging resistance.
- equivalents of SbPIN4 and SbPIN2 and other SbPINs are operative across other major cereal and crop species to enhance drought adaptation in localities worldwide where water limits crop growth post- anthesis.
- modulation of expression of a PIN selected from SbPINl to 1 1 such as SbPIN4 (Stgl) and SbPIN2 (Stg2) or their equivalent in another plant in all or selected tissue confers drought adaptation both directly, and indirectly, ultimately leading to higher grain yield, larger grain size, and lodging resistance under water-limited conditions.
- the stay-green phenotype involves the" presence of multiple proteins such as two or more of SbPINl to 1 1 such as SbPIN4 and SbPIN2.
- Taught herein is a method for generating a genetically modified plant which uses water more efficiently than a non-genetically modified plant of the same species, the method comprising introducing into a plant or parent of the plant a genetic agent which encodes two or more PINs or functional homolog or ortholog thereof which are associated with or facilitates a stay-green phenotype, which phenotype includes a shift in water use to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water limiting conditions; or an agent which modulates levels of expression of the two or more PINs.
- two or more means 2 or 3 or 4 or 5 or 6 or 7 or 8 or 9 or 10 or 1 1.
- a single or multiple PIN loci may also be expressed or inhibited.
- the genetic material is selected from (i) an agent which encodes SbPIN4; and (ii) an agent which modulates levels of expression of SbPIN4 or its equivalent in another plant.
- Increased water availability at anthesis is also achieved via increased water accessibility (better water extraction and deeper or greater lateral spread).
- T/LA transpiration per unit leaf area
- TE transpiration efficiency
- Increased TE via introgressing Stgl or Stg2 is proposed to be due to a) proportionally higher photosynthetic capacity compared with stomatal conductance, due to smaller, thinner and greener leaves, and/or b) a decrease in transpiration while maintaining biomass. Lysimetry studies indicate that both of these mechanisms contribute to higher TE in Stgl lines, with the reduction in transpiration the primary mechanism.
- Crop water use (CWU) before anthesis was negatively correlated with CWU after anthesis in an artificial dought (rain-out shelter [ROS]) experiment.
- ROS rain-out shelter
- StgX such as Stgl or Stg2 confers drought adaptation by being associated with pre- and post-anthesis biomass production.
- the Stgl or Stg2 region for example, reduces the pre:post anthesis biomass ratio below a critical level, increasing grain yield and lodging resistance.
- StgX such as Stgl, Stg2, Stg3 and/or Stg4 facilitates one or more of the following phenotypes:
- each of the key StgX mechanisms maps to a defined region, suggesting that the action of a single gene has multiple pleiotrophic effects.
- the present disclosure further teaches a business model to enhance economic returns from crop production.
- a business model for improved economic returns on crop yield comprising generating crop plants having a PIN expression profile resulting in the crop plant having a shift in water use by the plant to the post-anthesis period or increased accessibility of water during crop growth thereby increasing HI and grain yield under water-limited conditions, obtaining seed from the generated crop plant and distributing the seed to grain producers to yield enhanced yield and profit.
- Reference to a PIN includes SbPINl to 11 such as SbPIN4 and SbPIN2, as well as their equivalents in other plants.
- the plant management system includes the generation of drought adapted crop including cereal plants using the selection and modulated expression of PIN locus or its functional equivalent as herein defined alone or in combination with the introduction of other useful traits such as grain size, root size, salt tolerance, herbicide resistance, pest resistance and the like.
- the plant management system comprises generation of drought adapted plants and agricultural procedures such as irrigation, nutrient requirements, crop density and geometry, weed control, insect control, soil aeration, reduced tillage, raised beds and the like.
- the present specification is instructional as to a means to induce or enhance drought adaptation capacity in a plant by introducing do novo one or more features of the stay-green phenotype or elevating or reducing expression of an existing one or more PIN loci in a plant and/or selecting an PIN polymorphic variant with improved or enhanced expression or product activity.
- the manipulation of the stay-green phenotype may be done alone or as part of an integrated plant management system which may include further trait selection and/or improved agronomic techniques.
- the resulting crops use water more efficiently and have a higher yield of grain and increased grain size.
- the present teachings include business models to collect seed from drought adapted or enhanced crop plants for distribution to growers to ultimately increase grain yield.
- the present disclosure is enabling in the respect of the use of a genetic agent encoding a ⁇ protein or which modulates levels of expression of an indigenous PIN protein in the manufacture of a drought adapted plant.
- QTLs are identified herein as carrying one or more PIN loci wherein the level of expression of which is selected in breeding protocols or by genetic engineering, to promote a stay-green phenotype.
- Genetically modified plants and their progeny exhibiting the stay-green trait are also taught herein as well as seeds, fruit and flowers and other reproductive or propagating material.
- Genetic material which encodes a PIN protein or functional homolog or ortholog thereof which is associated with or facilitates a stay-green phenotype, which phenotype includes a canopy architecture which facilitates a shift in water use to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water limiting conditions; and an agent which modulates the level of the ⁇ ; are both enabled herein.
- the genetic material is selected from (i) an agent which encodes an SbPIN listed in Table 1A; and (ii) an agent which modulates levels of expression of an SbPIN listed in Table 1 A or its equivalent in another plant.
- the genetic material is selected from (i) an agent which encodes SbPIN4; and (ii) an agent which modulates levels of expression of SbPIN4 or its equivalent in another plant.
- the genetic material selected from (i) an agent which encodes SbPIN2; and (ii) an agent which modulates expression of SbPIN2 or its equivalent in another plant.
- the genetic material includes an agent which encodes a PIN, or functional homolog or ortholog thereof which is associated with or facilitates a stay-green phenotype, which phenotype includes a canopy architecture which facilitates a shift in water use to the post-anthesis period or increased accessibility of water during crop growth or increased transpiration efficiency resulting in increased harvest index and grain yield under water limiting conditions.
- the active agent includes an agent which up-regulates or down- regulating levels of the PIN.
- the subject genes may also be used as markers to transfer regions of genomes comprising one or more of the genes, or their equivalents in other plants, to a particular plant in order to include the stay-green phenotype.
- a quantitative trait locus which is referred to herein as Stgl which is an example of an StgX, has been identified which increases or enhances water use efficiency by sorghum plants.
- Stgl encodes a sorghum bicolor member of the auxin efflux carrier component 4 family, PIN4 (or SbPIN4).
- This major drought adaptation gene has been fine-mapped on the sorghum genome. Changes in auxin efflux could explain all of the multiple phenotypes observed in plants containing SbPIN4.
- the candidate gene (and promoter region) is sequenced in the two parents of the fine-mapping population (RTx7000 and Tx642) to identify a single nucleotide polymorphism.
- R A expression profiling of the Stgl fine-mapping population is also conducted for a subset of lines, times and organs. Phenotypes exhibited by SbPIN4 plants that could be explained directly by changes in auxin efflux include reduced tillering, smaller leaves (both length and width), reduced leaf mass and increased root:shoot ratio.
- Phenotypes exhibited by SbPIN4 plants that could be explained indirectly (or as emergent consequences of these direct effects) include increased availability of water at anthesis, higher leaf N concentration at anthesis, increased transpiration and biomass per unit leaf area, reduced pre:post anthesis biomass ratio, higher transpiration efficiency, retention of green leaf area during grain filling, increased harvest index, higher grain yield, larger grain size and increased lodging resistance. It is proposed that SbPIN4 works across other major cereal and crop species to enhance drought adaptation in localities worldwide where water limits crop growth post-anthesis.
- Stgl (SbPIN4) confers drought adaptation both directly, and indirectly, ultimately leading to higher grain yield, larger grain size, and lodging resistance under water-limited conditions.
- Increased water availability at anthesis is achieved via reduced water use due to two mechanisms (reduced tillering and smaller leaves) in plants containing the Stgl region. Both mechanisms, individually, appear to reduce canopy size by about 9%, on average.
- the 'low-tillering' mechanism dominates in low density environments when tillering potential is high.
- the 'small-leaf mechanism dominates in high density environments when tillering potential is low. Combined, these two mechanisms provide crop plants with considerable plasticity to modify canopy architecture in response to the severity of water limitation.
- Increased water availability at anthesis is also achieved via increased water accessibility (better water extraction and deeper or greater lateral spread).
- T/LA transpiration per unit leaf area
- TE transpiration efficiency
- Changes in transpiration per unit leaf area could be due to a) number of stomata, b) size of stomatal aperture, c) changes in the timing of stomatal opening and closing relative to VPD, and/or d) the number of hair base cells (which affects the boundary layer and hence T/LA).
- Introgressing Stgl into RTx7000 reduced the number of stomata and increased the number of hair base cells per unit leaf area in leaves 7 and 10; both mechanisms can conserve water by reducing T/LA.
- Introgressing Stgl into RTx7000 modified leaf anatomy by increasing the number of bundle sheath cells surrounding the vascular bundle.
- the increased number of cells in the bundle sheath might also contribute to increased photosynthetic assimilation and hence TE.
- Leaves 7 and 10 Differences in the morphology of leaves (e.g. Leaves 7 and 10) are apparent between Tx7000 and Stgl. In this case, there were more and smaller bundle sheaths surrounding the vascular bundle in Stgl . The increased number of cells in the bundle sheath might also contribute to increased photosynthetic assimilation and hence TE.
- Increased water use during grain filling is achieved via (i) increased water availability at anthesis and (ii) increased water accessibility (better water extraction and deeper or greater lateral spread) during grain filling. a) Increased water availability at anthesis
- Crop water use (CWU) before anthesis was negatively correlated with CWU after anthesis in an ROS experiment. For example, in one experiment a 25% increase in water use after anthesis (80 vs 60 mm) resulted in a 25% increase in grain yield (400 vs 300 g/m2). This translated to 50 kg/ha of grain for every additional mm of water available. b) Increased water accessibility during grain fitting
- Stgl region confers drought adaptation via a link between pre- and post-anthesis biomass production.
- the Stgl region reduces the pre:post anthesis biomass ratio below a critical level, increasing grain yield and lodging resistance.
- Each of the key Stgl mechanisms maps to the same region, indicating the action of a single gene with multiple pleiotrophic effects.
- RTx7000 produced 41% more (PO.05) culms/m2 than B35 (14.07 vs. 10.00).
- Introgression of the Stgl region alone into RTx7000 6078-1) reduced culms/m2 significantly (PO.05) compared with RTx7000 (9.40 vs. 14.07).
- additional introgressions of either Stg2 or Stg4 increased culm numbers to 10.49 (1,2 combination) and 10.74 (1,4 combination).
- RTx7000 produced almost eight-fold more (PO.05) GLAAt than B35 (15460 vs. 1980). Introgression of the Stgl region alone into RTx7000 (6078-1) reduced GLAAt significantly (PO.05) compared with RTx7000 (3121 vs 15460). Compared with Stgl only, additional introgressions of either Stg2 or Stg4 increased GLAAt to 4187 (1,2 combination) and 4797 (1,4 combination). All lines containing Stgl (in any combination) were not significantly different (PO.05) in GLAAt from Stgl alone.
- a Stgl fine-mapping population was grown in the field under high and low density conditions in three consecutive years. The number of culms per plant was measured at anthesis in each year and a combined analysis was undertaken across years. Overall, RT 7000 produced 47% more culms per plant than 6078-1 (1.85 vs. 1.26; Figure 3). [00175] In these field studies, the trait (culm number per plant) was mapped. An arbitrary value of 1.54 culms per plant gave the optimal separation between high and low tillering for mapping purposes (i.e. recombinants with less than 1.54 culms were BB genotypes while those with more than 1.54 culms were TT genotypes).
- Stepping down through the markers reveals that gain-of-function (low tillering) was achieved in three genotypes (10564-2, 10704-1 and 10620-4).
- One recombinant (10568-2) exhibited a high tillering phenotype, while three others (10620-4, 10704-1 and 10564-2) exhibited low tillering phenotypes.
- An auxin efflux carrier component 5 gene was proposed to be the candidate gene.
- a subset of the Stgl fine-mapping population was grown in the field at the Rain- Out Shelter (ROS) under high and low water conditions, with each water treatment split for high and low density. This created four water regimes with increasing levels of water deficit: HWLD (least stressed) ⁇ HWHD ⁇ LWLD ⁇ LWHD (most stressed).
- HWLD low Density
- RTx7000, 10568-2 and 10709-5 produced 27% more culms per plant than 6078-1 and 10604-5 under LWLD conditions (2.05 vs. 1.62; Figure 4), and 23% more culms per plant under HWLD conditions (1.49 vs. 1.22).
- T4 tiller numbers also varied among genotypes. 6078-1 produced a T4 tiller in 3 of 4 replicates, while RTx7000 produced a T4 in all 4 replicates. Hence the Stgl introgression essentially prevented the growth of T2 and T3 tillers in a RTx7000 background.
- Table 4 shows the presence of tillers (T1 -T3) and total tiller number for eight high-tillering recombinants (brown shading) and eight low-tillering recombinants (green shading) from the Stgl fine-mapping population.
- Gain-of-function is achieved in recombinant 10604-1-157-5. Gain- of-function (low tillering) is also achieved in three recombinants (10604-1-195-5, 10604-1- 56-7, 10604-1-477-4).
- Table 5 shows the presence of tillers (T1-T4), including secondary tillers, and total tiller number for five high-tillering recombinants (brown shading) and three low- tillering recombinants (green shading) from the Stgl fine-mapping population.
- Stgl confers two mechanisms for reducing canopy size: a) reduced tillering, and b) reduced leaf size. Combined, these two mechanisms provide a fair degree of plasticity for the plant to modify canopy architecture in response to environmental and/or management factors.
- the 'tails' of the Stgl fine-mapping population were selected. Two genotypes exhibited particularly long leaves (10604-1-157-5 and 10604-1 - 318-1) and three genotypes exhibited particularly short leaves (10604-1-222-1, 10604-1- 501-327-3 and 6078-1).
- Gain-of-function (short leaf) is achieved in recombinant 10604-1-222-1.
- the 'small leaf gene is proposed to map to the same region as the Mow tillering' gene.
- the 'tails' of the Stgl fine-mapping population were selected ( Figure 16). Three genotypes exhibited particularly long leaves (10604-1-157-5, 10604-1-318-1 and RTx7000) and two genotypes exhibited particularly short leaves ( 10604- 1 -222- 1 and 6078- 1 ).
- Reduced crop water use at anthesis can be caused by a) a smaller canopy size with equivalent transpiration per unit leaf area, b) an equivalent canopy size with lower transpiration per unit leaf area, or c) a smaller canopy size and lower transpiration per unit leaf area.
- Transpiration is the product of leaf area (LA) and transpiration per leaf area (T/LA).
- LA was similar between Stgl and RTx7000 (11795 vs 1 1628 cm2), yet T/LA was less in Stgl than Tx7000 (2.60 vs 2.85), resulting in less water use per plant (T) in Stgl than RTx7000 (30.7 vs 32.8 1).
- water savings in Stgl in a high VPD environment
- were achieved entirely by a reduction in T/LA suggesting that this is a constitutive water conservation strategy conferred by Stgl.
- higher transpiration efficiency (TE) in Stgl was a consequence of equivalent biomass and lower transpiration.
- T Transpiration
- RTx7000 Transpiration (T) is the product of LA and T/LA.
- LA was 31% less in Stgl than RTx7000 (4898 vs 7082 cm2). This was offset slightly by a 9% increase in T/LA in Stgl compared with RTx7000 (5.15 vs 4.70).
- the net result was a 22% reduction in water use per plant (T) in Stgl compared with RTx7000 (25.6 vs 32.7 1), primarily due to reduced canopy size.
- the increase in T/LA exhibited by Stgl may itself be a drought adaptation mechanism, cooling the leaf and enabling photosynthesis to continue.
- the plasticity in T/LA appears to be particularly important in the regulation of plant water status.
- Increased water availability at an thesis may also be achieved via increased water accessibility due to better water extraction and or deeper or greater lateral spread of roots in plants containing the Stgl region
- Root mass and root: shoot ratio were higher in Stgl than RTx7000 at the Leaf 6 stage. There was considerable transgressive segregation for these traits in the Stgl fine-mapping population. The relation between root mass and roo shoot ratio highlights the opportunity for further genetic advance in these traits.
- Root mass per leaf area ratio can be used as a drought adaptation index at the seedling stage since it integrates the capacity of the plant to access water (root mass) with the capacity of the plant to utilise water (leaf area). A higher index indicates a greater capacity to access water per unit leaf area. Stgl exhibited a higher root mass per leaf area ratio relative to RTx7000 due to both a higher root mass and a smaller leaf area.
- Table 7 shows mainstem, tiller and total biomass per leaf area for RTx7000 (recurrent parent) and a number of near-isogenic lines containing various Stgl introgressions grown under high and low water stress at Biloela, Queensland, Australia.
- Stgl and RTx7000 displayed equivalent B/LA under low water stress. However, T was ⁇ 7% lower. in Stgl, due to ⁇ 10% lower T/LA which, in turn, increased TE by ⁇ 9% ( Figure 45). Therefore, Stgl maintained biomass but used less water compared with RTx7000.
- B/LA was -6% higher in Stgl compared with RTx7000.
- B/LA was positively correlated with T/LA but not with TE.
- the higher B/LA displayed by Stgl was due to higher T/LA.
- B/LA was positively correlated with T/LA and negatively correlated with TE under high water stress.
- Stgl used ⁇ 22% less water than RTx7000 during the pre-anthesis period. Therefore, Stgl would have significantly more water available to fill grain, despite lower biomass at anthesis.
- Increased TE via introgressing Stgl may be due to a) proportionally higher photosynthetic capacity compared with stomatal conductance, due to smaller, thinner and greener leaves, or b) a decrease in transpiration per leaf area while maintaining biomass per leaf area
- Greener leaves may increase photosynthetic capacity and therefore water use efficiency.
- photosynthesis increased with SPAD value until reaching a plateau at a SPAD of -48.5 ( Figure 26).
- the line (6078-1) with the highest SPAD value (51.6) exhibited a relatively low rate of photosynthesis (32.1 MJ/m2/d). This result is either a) anomalous, or b) indicates a real decline in photosynthesis at high SPAD values.
- Leaf greenness (SPAD) and WUE were positively correlated in a subset of the Stgl fine-mapping population (Figure 27).
- Increased TE via introgressing Stgl may be due to a) proportionally higher photosynthetic capacity compared with stomatal conductance, due to smaller, thinner and greener leaves, or b) a decrease in transpiration per leaf area while maintaining biomass per leaf area
- Transpiration efficiency was negatively correlated with transpiration per leaf area (T/LA) under low and high VPD conditions ( Figure 29) in a set of Stg NILs, including the recurrent parent (RTx7000).
- RTx7000 transpiration per leaf area
- Figure 29 the ranking of Stg NILs relative to RTx7000 interacted with VPD conditions.
- T/LA in Stgl was lower relative to RTx7000 under high VPD conditions, yet higher than RTx7000 under low VPD conditions.
- Changes in transpiration per unit leaf area could be due to a) number of stomata, b) stomatal aperture, c) changes in the timing of stomatal opening and closing relative to VPD, and/or d) numbe of hair base cells (which affects the boundary layer and hence
- T/LA Introgressing Stgl into RTx7000 variously affected T/LA, depending on VPD conditions. Relative to RTx7000, Stgl increased T/LA by ⁇ 9% under low VPD and decreased T/LA by -10% under high VPD.
- T/LA can be regulated by a) the number of stomata per unit leaf area, b) the size to the stomatal aperture, c) the timing of stomatal opening and closing, and/or d) the number of hair base cells (which affects the boundary layer and hence T/LA). Measurements of two of these four components (a and d) have been made.
- Increased water use during grain filling is achieved via (i) increased water availability at anthesis and (ii) increased water accessibility (better water extraction and deeper or greater lateral spread) during grain filling a) Increased water availability at anthesis
- Crop water use (CWU) before anthesis was negatively correlated with CWU after anthesis in the ROS experiment ( Figure 30). For example, saving 20 mm of water before anthesis (165 vs 185 mm) enabled the utilization of an additional 20 mm after anthesis (80 mm vs 60 mm). So all of the water conserved before anthesis was utilized by the crop after anthesis. Overall, a 25% increase in water use after anthesis in this experiment resulted in a 25% increase in grain yield (400 vs 300 g/m2). This translated to 50 kg/ha of grain for every additional mm of water available. While these data support the concept that increased water use during grain filling is achieved via increased water availability at anthesis, it does not explain about increased water accessibility during grain filling. b) Increased water accessibility during grain filling
- Post-anthesis stem mass is a component of lodging resistance. Analysis of this component provides some understanding of how Stg introgressions affect lodging resistance. Reducing PPBR from >8 to ⁇ 4 resulted in a gradual increase in post-anthesis stem mass. However, further reducing PPBR below 4 resulted in a relatively sharp increase in post-anthesis stem mass. Introgressing Stgl into RTx7000 increased post-anthesis stem mass under both LD (marginal increase) and HD (significant increase) conditions.
- Canopy size as evidenced by GLAA, largely determined the ratio of pre:post anthesis biomass (Figure 34). Under both high and low density treatments, introgressing Stgl into a RTx7000 background reduced GLAA which, in turn, reduced the ratio of pre:post anthesis biomass to ⁇ 3, ensuring adequate water availability for grain filling under these water-limited conditions.
- post anthesis stem mass (PASM) is used as a surrogate for lodging resistance.
- PPBR post anthesis stem mass
- a high pre:post anthesis biomass ratio increased the amount of stem reserves remobilized during grain filling, thus reducing stem biomass and increasing the likelihood of lodging.
- Stgl a significantly reduced the amount of stem reserves mobilized under LD ( ⁇ 5 vs 65 g/m2) and HD (-80 vs 140 g/m2).
- Stgl significantly reduced the amount of stem reserves mobilized under HD (-80 vs 140 g/m2), but not LD.
- stem reserves mobilized was greater under HD than LD, reflecting the greater water deficit under HD.
- the difference in stem reserve mobilization between HD and LD was twofold in RTx7000 (about 140 vs 70 g/m2).
- Grain yield remained low (at a benchmark of -4.2 t/ha) until the pre:post anthesis biomass ratio fell below -3 (HD) or -2.5 (LD) [Figure 38].
- grain yield increased significantly for each incremental reduction in these ratios, with the rate of increase in grain yield being slightly higher under LD than HD. This suggests that post-anthesis water availability was closely linked to pre-anthesis GLAA and biomass, and that a certain reduction in GLAA was required to ensure adequate water availability for grain filling.
- Stgl reduced the pre:post anthesis biomass ratio below the critical levels, resulting in yield increases of 28% (LD) and 22% (HD), relative to RTx7000.
- pre-anthesis biomass varied by only -5% (from 522 to 552 g m2) among genotypes, yet post-anthesis biomass varied almost twofold (from 173 to 313 g/m2).
- post-anthesis biomass varied almost twofold (from 173 to 313 g/m2).
- 10709-5 and RTx7000 both produced -550 g/m2 of pre- anthesis biomass, yet the Stgl recombinant (10709-5) produced -60% more post-anthesis biomass.
- PAB post-anthesis biomass
- PAB increased for each incremental reduction in CWU down to a level of 175 mm, with PAB plateauing at about 310 g/m.2. Further reductions in CWU at anthesis below 175 mm did not result in additional PAB.
- Canopy size as evidenced by GLAA, largely determined the ratio of pre:post anthesis biomass (PPBR).
- PPBR pre:post anthesis biomass
- introgressing Stgl (or particular recombinants such as 10709-5) into a RTx7000 background reduced GLAA which, in turn, reduced the ratio of pre:post anthesis biomass, thereby increasing water availability for grain filling under these water-limited conditions.
- the PPBR value for 6078-1 appears anomalous (too high) since this genotype is placed well above the GLAA/PPBR regression line.
- a high pre:post anthesis biomass ratio increased the amount of stem reserves remobilized during grain filling, thus reducing stem biomass and increasing the likelihood of lodging.
- Stg 1 significantly reduced the amount of stem reserves mobilized under HD (-100 vs 160 g/m2).
- the extent of stem reserves mobilized was greater under HD than LD, reflecting the greater water deficit under HD.
- the difference in stem reserve mobilization between HD and LD was more than twofold in RTx7000 (about 160 vs 60 g m2).
- PASM increased with decreasing PPBR over the whole range of PPBR (1.5 - 6), whereas grain yield, and to a lesser extent PAB, only increased when PPBR fell below -3. This suggests that relatively small water savings before anthesis were still able to improve lodging resistance, although greater water savings were required before grain yield responded.
- CWU during grain filling remained low (at a benchmark of -60 mm) until the pre:post anthesis biomass ratio fell below -3.5 (Figure 39). Below this critical value, CWU during grain filling increased significantly for each incremental reduction in this ratio. Since none of the Stgl introgressions reduced the PPBR to ⁇ 3.5 under HD, no increases in CWU during grain filling were realised from Stgl recombinants in this treatment. Under LD, some of the Stgl introgressions (and RTx7000) reduced PPBR below the critical level, increasing CWU during grain filling up to ⁇ 80 mm compared with the HD baseline ( ⁇ 60 mm).
- Delayed leaf senescence (stay-green) is a consequence of higher plant water status during grain filling (due to increased water use)
- Plant water status was determined on FL-2 (two leaves below the flag) at mid- grain filling using two methodologies: leaf water potential (LWP) and relative water content (RWC).
- LWP leaf water potential
- RWC relative water content
- LWP was measured in the field with a pressure bomb. Following determination of LWP in the field, a sample of the same leaf was placed on ice and, within a few minutes, taken to a laboratory some 300 m away for determination of RWC by standard methods.
- the RWC of FL-2 was negatively correlated with the relative rate of leaf senescence at mid-grain filling under both high and low densities in a set of Stg NILs including the recurrent parent.
- Increased lodging resistance is a consequence of higher plant water status during grain filling (due to increased water use)
- Post-anthesis biomass is mainly comprised of a) post-anthesis stem mass (PASM), a measure of stem reserve mobilization and a component of lodging resistance, and b) grain yield.
- PASM post-anthesis stem mass
- Grain-growers require that both grain yield and lodging resistance be maximized, i.e. they do not want one at the expense of the other.
- Post-anthesis stem mass was highly linearly correlated with PAB under HD and LD conditions ( Figure 44). While Stgl had little impact on PASM under the milder drought (LD), the amount of dry mass translocated from the stem during grain filling was much less in Stgl compared with RTx7000 (85 vs. 139 g/m2) under the more severe drought (HD). This resulted in greater stem mass at maturity in Stgl relative to RTx7000 (286 vs. 204 g/m2) which, in turn, should have increased lodging resistance.
- LD milder drought
- Post-anthesis stem mass was highly linearly correlated with PAB under HD and LD conditions ( Figure 46).
- the HD and LD correlations were almost parallel, although offset by about 50 g/m2, i.e. for a given level of PAB, say 300 g m2, PASM was -50 g/m2 less in HD than LD (-100 vs. -50 g m2). This reflects the higher level of stress in the HD treatment.
- both Stgl and Stg la utilized -80 g/m2 of stem reserves compared with almost 140 g/m2 in RTx7000, yet Stgl produced -28% more PAB than Stgl a for equivalent stem reserve utilisation.
- Stgl a and to a lesser extent Stgl both increased PAB relative to RTx7000.
- Grain yield was positively correlated with PAB under HD and LD.
- HD Stgl outyielded RTx7000 by 24% although Stg la was equivalent to RTx7000 in grain yield.
- LD Stgl and Stg la outyielded RTx7000 by 42% and 20%, respectively.
- Stgl utilized more stem reserves compared with RTx7000 (87 vs 66 g/m2). Overall, Stgl increased stem mass at maturity by 22% (HD) and 16% (LD) relative to RTx7000. Also, Stg la utilized significantly less stem reserves than RTx7000 under both crop densities.
- Post-anthesis stem mass was highly linearly correlated with PAB under HD and LD conditions (Figure 48). Genetic variation in utilization of post-anthesis stem reserves ranged from -30-110 g/m2 under LD, and from -100-160 g/m2 under HD, reflecting the higher level of stress in the HD treatment. Under HD, the Stgl parent (6078- 1) and two of the Stgl recombinants (10604-5 and 10709-5) utilized significantly less stem reserves compared with RTx7000 (-100 vs.160 g/m2), yet produced more PAB than RTx7000 (-170 vs. 130 g/m2).
- RWC at mid-grain filling in FL-2 was positively correlated with grain yield under HD and LD.
- grain yield was higher under LD than HD for a given level of RWC.
- RWC and grain yield were higher in Stgl than RTx7000 under both crop densities. For example in Stgl under HD, a 26% increase in RWC was associated with a 58% increase in grain yield, relative to RTx7000.
- C WU during grain filling remained low (at a benchmark of ⁇ 58 mm for HD) until the pre.post anthesis biomass ratio fell below -3.5.
- the PPBR did not drop below the critical threshold in any genotype under HD, hence C WU during grain filling remained relatively low for all genotypes in this treatment.
- CWU during grain filling increased significantly for each incremental reduction in this ratio. Only one Stgl recombinant (10709-5) increased CWU during grain filling relative to RTx7000.
- the score (S value: a measure of the similarity of the query to the sequence shown), E-value (the probability due to chance, that there is another alignment with a similarity greater than the given S score), %ID and length of sequence homology for each of the 1200 hits were collated.
- the relationship between the 4 measures was analyzed and the S score was selected as the main measure to assess likelihood of sequence similarity.
- 3 S score categories were identified (>1000; >499 and ⁇ 1000; ⁇ 499) and a list of 1 1 sorghum genes with scores >499 (i.e. in the first 2 categories) was produced (Table 8).
- the aim of the experiment was to measure expression levels of genes that are identified herein as stay-green gene candidates under well-watered conditions and after a drought stress has been imposed on the plants to see whether there were any differences in expression in the stay-green compared with the senescent plants.
- the experiment was divided into two parts: an early drought stress (Expl) and a late drought stress (Exp2).
- SbPIN4 (Stg 1 WW: low in leaves, Lower expression WW: Higher in Tx7000 vs candidate) high in roots in most tissues BTx642 and Stgl in roots
- SbPIN2 WW high in leaves, Increase in roots, WW: Higher in Stg2/BTx642
- SbPIN4 was generally (across all conditions) more highly expressed in roots and stems and less expressed in leaves, while SbPIN2 generally showed higher expression in leaves and stems and lower expression in roots).
- aspects described herein are susceptible to variations and modifications other than those specifically described. It is to be understood that those aspects include all such variations and modifications. Aspects herein described also include all of the steps, features, compositions and compounds referred to or indicated in this specification, individually or collectively, and any and all combinations of any two or more of the steps or features. [00311] Those skilled in the art will appreciate that aspects described herein are susceptible to variations and modifications other than those specifically described. It is to be understood that these aspects include all such variations and modifications. Such aspects also include all of the steps, features, compositions and compounds referred to or indicated in this specification, individually or collectively, and any and all combinations of any two or more of the steps or features. 11 001478
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MX2013005485A MX352184B (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants. |
US13/885,357 US20130333066A1 (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
EP11841328.5A EP2640182B1 (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
CN201180062760XA CN103429072A (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
EP18180109.3A EP3434099A1 (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
EA201390724A EA034044B1 (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
BR112013012025A BR112013012025A2 (en) | 2010-11-15 | 2011-11-15 | drought tolerant plants |
UAA201307591A UA113503C2 (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
CA2817758A CA2817758C (en) | 2010-11-15 | 2011-11-15 | Methods for generating drought tolerant plants |
AU2011331908A AU2011331908B2 (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
ES11841328.5T ES2692857T3 (en) | 2010-11-15 | 2011-11-15 | Plants tolerant to drought |
PL11841328T PL2640182T3 (en) | 2010-11-15 | 2011-11-15 | Drought tolerant plants |
ZA2013/03689A ZA201303689B (en) | 2010-11-15 | 2013-05-21 | Drought tolerant plants |
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Cited By (3)
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WO2013071366A1 (en) * | 2011-11-16 | 2013-05-23 | The State Of Queensland Acting Through The Department Of Agriculture, Fisheries And Forestry | Drought tolerant plants produced by modification of the stay-green stgx locus |
CN107988236A (en) * | 2017-12-13 | 2018-05-04 | 南京农业大学 | The genetic engineering application of auxin transport protein gene of paddy rice OsPIN9 |
CN112430604A (en) * | 2020-12-11 | 2021-03-02 | 河南农业大学 | Genetic engineering application of gene OsPIN10b |
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WO1997013843A1 (en) * | 1995-10-12 | 1997-04-17 | Cornell Research Foundation, Inc. | Production of water stress or salt stress tolerant transgenic cereal plants |
EP2055779A3 (en) * | 1999-05-07 | 2009-08-12 | E.I. Dupont De Nemours And Company | Auxin transport proteins |
AU2002341541A1 (en) * | 2001-06-22 | 2003-03-03 | Syngenta Participations Ag | Abiotic stress responsive polynucleotides and polypeptides |
US7256327B2 (en) * | 2002-11-29 | 2007-08-14 | The University Of Hong Kong | Genetically modified plants expressing proteinase inhibitors, SaPIN2a or SaPIN2b, and methods of use thereof for the inhibition of trypsin- and chymotrypsin-like activities |
CN1813060A (en) * | 2003-04-15 | 2006-08-02 | 巴斯福植物科学有限公司 | Plant cells and plants with increased tolerance to environmental stress |
CN100362104C (en) * | 2004-12-21 | 2008-01-16 | 华中农业大学 | Using gene of transcriptional factor OSNACX of paddy to increase drought resistance and salt tolerant abilities of plants |
CN101228279A (en) * | 2005-07-19 | 2008-07-23 | 巴斯福植物科学有限公司 | Yield increase in plants overexpressing the MTP genes |
US8362325B2 (en) * | 2007-10-03 | 2013-01-29 | Ceres, Inc. | Nucleotide sequences and corresponding polypeptides conferring modulated plant characteristics |
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Cited By (6)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
WO2013071366A1 (en) * | 2011-11-16 | 2013-05-23 | The State Of Queensland Acting Through The Department Of Agriculture, Fisheries And Forestry | Drought tolerant plants produced by modification of the stay-green stgx locus |
EA030382B1 (en) * | 2011-11-16 | 2018-07-31 | Дзе Стейт Оф Квинсленд Эктинг Тру Дзе Департмент Оф Эгрикалчер Энд Фишериз | DROUGHT TOLERANT PLANTS PRODUCED BY MODIFICATION OF THE STAY-GREEN StgX LOCUS |
US10590431B2 (en) | 2011-11-16 | 2020-03-17 | The State Of Queensland | Drought tolerant plants produced by modification of the stay-green STGX locus |
CN107988236A (en) * | 2017-12-13 | 2018-05-04 | 南京农业大学 | The genetic engineering application of auxin transport protein gene of paddy rice OsPIN9 |
CN107988236B (en) * | 2017-12-13 | 2021-02-09 | 南京农业大学 | Genetic engineering application of oryza sativa auxin transport protein gene OsPIN9 |
CN112430604A (en) * | 2020-12-11 | 2021-03-02 | 河南农业大学 | Genetic engineering application of gene OsPIN10b |
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CN108624618A (en) | 2018-10-09 |
AU2011331908A1 (en) | 2013-04-18 |
CN103429072A (en) | 2013-12-04 |
EP2640182A1 (en) | 2013-09-25 |
TR201815884T4 (en) | 2018-11-21 |
PL2640182T3 (en) | 2019-02-28 |
HUE040165T2 (en) | 2019-02-28 |
US20130333066A1 (en) | 2013-12-12 |
EA034044B1 (en) | 2019-12-23 |
EP2640182B1 (en) | 2018-07-25 |
ZA201303689B (en) | 2014-07-30 |
EA201390724A1 (en) | 2014-05-30 |
EP2640182A4 (en) | 2014-05-07 |
CA2817758C (en) | 2019-01-29 |
ES2692857T3 (en) | 2018-12-05 |
EP3434099A1 (en) | 2019-01-30 |
CA2817758A1 (en) | 2012-04-24 |
MX2013005485A (en) | 2013-09-26 |
UA113503C2 (en) | 2017-02-10 |
MX352184B (en) | 2017-11-09 |
AU2011331908B2 (en) | 2015-05-07 |
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