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11. Wearing sterile latex gloves, a mask and a watchmaker's 2x headband enlarger, excise the haploid embryos from the disinfected GPCs under a laminar flow hood. (Fine scalpels fashioned from discarded dentist's needles are best for this purpose; their surgical steel enables repeated flaming, and cooling in cold sterile water. Lift each green caryopsis from the McCartney bottle by inserting the sterilized scalpel into its attachment end. Hold the brush end between the thumb and forefinger and cut along the lateral grooves at the attachment end, fold back the dorsal pericarp and green integuments, and lift out the haploid embryo).

12. Five excised embryos are placed in each 100 ml tissue culture jar containing 20 ml of autoclaved IRM (the barley anther culture medium of Olsen (1987) modified by Daniel (1990) and us. This modified MS-medium contains 165 mg
l-1 instead of 1650 mg l-1 NH4NO3, 20000 mg l-1 sucrose, 100 mg l-1 myoinositol, 0.4 mg l-1 thiamine HCl, 0.5 mg l-1 BA, 1 mg l-1 IBA and 2300 mg l-1 Gelrite; the pH is adjusted to 5.8 with KOH).

13. Seal the transparent lids of the tissue culture jars containing the rescued embryos with clear plastic wrap. Place the jars in the dark for 10 days at 6-8C and then at 25C until the regenerating shoots are 1 cm long. Transfer them to a 25C growth cabinet with cool white fluorescent tubes on a 10 h light/14 h dark cycle. The regenerating plantlets reach the three leaf stage after four to six weeks.

14. Transplant the plantlets in micro-irrigated, sand-filled 2-liter pots (6-8 per pot) in a 16C growthroom on a 10 h light/14 h dark cycle, and water once with a dilute seaweed extract (2 ml/l Kelpak) to stimulate growth. (Plantlets transplanted in a greenhouse should be covered with a 50% shade screen for a week).

15. Lift the plants from the pots as soon as they have developed two to three sprouts (usually after a month), and trim their washed roots to 10 em. Treat the plants in an aerated (use an aquarium pump) aqueous solution of 0.05% colchicine (BDH), without DMSO, for 24 hours at 22C. Rinse the plants in running water for an hour and transfer them to aerated water in a refrigerator at 4C for three days. (The colchicine solution can be used again; filter and store in a refrigerator).

16. The shoots of the treated plants are cut back to 15 cm and the plants repotted (2-3 per pot) in the 16C growthroom. Water once with the Kelpak solution.

17. After 4-6 weeks the + or - 95% plants that survived the colchicine treatment will have formed new shoots. Transfer the potted spring wheats to a greenhouse on a 18C day/12C night cycle, but vernalize the winter wheats first for at least six weeks at 4C.

18. Nearly all the colchicine treated plants will produce a few spikes with fertile doubled haploid (DH) sectors. Cover the fertile spikes to prevent cross pollination. These spikes will give rise to homozygous DH lines from which new cultivars can be selected.


References

Bajaj YPS (ed) (1990) Biotechnology in agriculture and forestry 12. Haploids in crop improvement I. Springer Verlag, Berlin.

Barclay IR(1975)
High frequency of haploid production in wheat (Triticum aestivum) by chromosome elimination. Nature 256: 410-411.

Daniel G
(1990) Einfluss verschiedener Faktoren auf die Pflanzenregeneration in der Antherenkultur bei@Sommerund Wintergerste. Bayer Landw Jahrbuch 67: 609-617.

Henry Y and De Buyser J (1990)
Wheat anther culture. In: Bajaj YPS (ed) Biotechnology in agriculture and forestry 13, Wheat. Springer-Verlag, Berlin, 285-352.

Inagaki MN and Tahir M
(1990) Comparison of haploid production frequencies in wheat varieties crossed with Hordeum bulbosum L. and maize. Jpn J Breed 40:209-216.

Jensen CJ
(1977) Monoploid production by chromosome elimination. In: Reinert J and Bajaj YPS (eds) Applied and fundamental aspects of plant cell, tissue and organ culture. Springer Verlag, Berlin, 299-340.

Kisana NS, Nkongolo KK, Quick JS and Johnson DL (1993) Production of doubled haploids by anther culture and wheat x maize method in a wheat breeding programme. Plant Breed 110: 96-102.

Laurie DA and Bennett MD (1988)
The production of haploid wheat plants from wheat x maize crosses. Theor Appl. Genet 76: 393-397.

Laurie DA and Reymondie S (1991)
High frequencies of fertilization and haploid seedling production in crosses between commercial hexaploid wheat varieties and maize. Plant Breed 106: 182-189.

Matzk F and Mahn A (1994)
Improved techniques for haploid production in wheat using chromosome elimination. Plant Breed 113: 125-129.

Mujeeb-Kazi A, Riera-Lizarazu 0 and William MDHM
(1995) Production of polyhaploid wheat plants using maize and Tripsacum. CIMMYT Research Report 2: 47-65.

Olsen, FL
(1987) Induction of microspore embryogenesis in cultured anthers of Hordeum vulgare. The effects of ammonium nitrate, glutamine and asparagine as nitrogen sources. Carlsberg Res Commun 53: 393-404.

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