Hostas Analyzed by Flow Cytometry

Scientists can measure the amount of DNA within the nucleus of a plant cell by using a technique known as flow cytometry. This, in turn, is helpful to determine the ploidy of hosta cultivars and species. In addition, DNA testing can identify ploidy chimeras that have different ploidy among the three apical layers found within the meristem of a hosta. (The three apical layers of cells within the meristem, or center, of a hosta can be genetically different. These layers are commonly designated as L1, L2, and L3.)

Like most plants, hostas generally are diploid, which means they have two sets of chromosomes. On the other hand, a plant may contain four sets of chromosome in each apical layer and therefore be a tetraploid (4-4-4). With a ploidy chimera, not all of the apical layers have the same ploidy. An example of a ploidy chimera is 'Grand Tiara' (4-2-2) which has four sets of chromosomes in layer 1 and two sets of chromosomes in layers 2 and 3.

Occasionally, a plant may have an abnormal number of chromosomes in one or more of the apical layers. This is a case in which there are extra or missing chromosomes and is known as aneuploidy. An example of an aneuploid hosta is 'Mango Tango' (4-2.8-4) which has an abnormal number of chromosomes in the center apical layer of the meristem. For the aneuploid hostas which have been tested, the ploidy of each layer is given when it can be determined.

In the case of 'Sum and Substance', seedlings from open pollination show a loss of DNA compared with the mother plant. However, the ploidy of these seedlings cannot be calculated when the pollen parent is unknown. The contribution of 'Sum and Substance', being triploid, to the seedlings is unknown. Therefore, when the pollen parent is unknown too, there is no frame of reference to calculate the ploidy of the seedlings. In such cases, the ploidy status is simply listed as "Aneuploid".

The flow cytometry measurements were conducted by Ben J. M. Zonneveld, formally at Leiden University in the Netherlands and currently at Naturalis Biodiversity Center in Leiden.

  • 'Abiqua Drinking Gourd' 2-2-2
  • 'Academy Ametrine' Aneuploid
  • 'After Midnight' 3.8-2-2
  • 'Aki Tenryu' 2-2-2
  • 'Allegan Fog' 2-2-2
  • 'Amanuma' 2-2-2
  • 'Amber Tiara' 4-4-4
  • 'Amine Tachi' 3-3-3
  • 'Aphrodite' 2-2-2
  • 'Atlantis' 4-2-2
  • 'Avocado' 4-2-2
  • 'Barney Fife' 2-2-2
  • 'Beatrice' 2-2-2
  • 'Bedazzled' 2-2-2
  • 'Bella' 2-2-2
  • 'Belle of the Ball' 2-2-2
  • 'Bill Brincka' 2-2-2
  • 'Blauwichtel' 2-2-2
  • 'Blazing Saddles' 3-3-3
  • 'Blizzard' 3-3-3
  • 'Blue Cadet' 2-2-2
  • 'Blue Ivory' 4-2-2
  • 'Blue Mouse Ears' 2-2-2
  • 'Blue Splendor' 2-2-2
  • 'Blue Umbrellas' 2-2-2
  • 'Blythe Spirit' 2-2-2
  • 'Botanical Look' 2-2-2
  • 'Bottom Line' 3-3-3
  • 'Breathless' 2-2-2
  • 'Bubba' Aneuploid (see footnote 1)
  • 'Bulletproof' 4-4-4
  • 'Bunter Babette' 2-2-2
  • 'Buster'
    (1) 3.4-3.4-3.4
    (2) 3.5-3.5-3.5
  • 'Butter Rim' 2-2-2
  • 'Cadillac' 2-2-2
  • capitata 2-2-2
  • 'Captain Kirk' 4-2-2
  • 'Carolina Sunshine' 2-2-2
  • 'Cascades' 2-2-2
  • 'Cerveza' 2-2-2
  • 'Cheatin Heart'
    (1) 2-2-2
    (2) 2-2-4 (see footnote 2)
  • 'Chelsea Ore' 2-2-2
  • 'Cherry Berry' 2-2-2
  • 'Cherry Tomato' 2-2-2
  • 'Chinese Sunrise' 2-2-2
  • 'Chionea' 2-2-2
  • 'Chishima Iwa' 2-2-2
  • 'Choko Nishiki' 2-2-2
  • 'Christmas Candy' 4-4-4
  • clausa var. clausa 3-3-3
  • clavata 2-2-2
  • 'Climax' 2-2-2
  • 'Collector's Choice' 2-2-2
  • 'Concord' 2-2-2
  • 'Cool as a Cucumber' 2-2-2
  • 'Corona' 3-3-3
  • 'County Park' 2-2-2
  • 'Crispula' 2-2-2
  • 'Crystal Chimes' 2-2-2
  • 'Cynthia' 2-2-2
  • 'Dance with Me' 4-4-4
  • 'Dancing in the Rain' 2-2-2
  • 'Dancing Stars' 2-2-2
  • 'David A. Haskell' 3-3-3
  • 'Decorata' 2-2-2
  • 'Dee's Golden Jewel' 2-2-2
  • 'Devon Tor' 2-2-2
  • 'Diamond Tiara' 2-2-2
  • 'Diana Remembered' 2-2-2
  • 'Domain de Courson' 3-3-3
  • 'Dorset Blue' 2-2-2
  • 'Doubled Up' 4-4-4
  • 'Dream Weaver' 4-2-2
  • 'Dress Blues' 2-2-2
  • 'Dust Devil' 4-4-4
  • 'Eagle's Nest' 3-3-A
  • 'El Niño' 2-2-2
  • 'Elata' 2-2-2
  • 'Elatior' 2-2-2
  • 'Elegans' 2-2-2
  • 'Embroidery' 2-2-2
  • 'Emerald Necklace' 2-2-2
  • 'Eternal Flame' 4-4-4
  • 'Enterprise' 4-2-2
  • 'Faithful Heart' 2-2-2
  • 'Fall Bouquet' 2-2-2
  • 'Fantabulous' 2-2-2
  • 'Filagree' 2-2-2
  • 'Final Summation' 2.9-2.9-2.9
  • 'Final Victory' 4-4-4
  • 'Fire and Ice' 4-4-4
  • "Fire Works"(see footnote 3)
  • 'Fireworks' 3.2-4-4
  • 'First Light' 4-4-6
  • 'Five O'Clock Shadow'
    (1) 2-2-2
    (2) 2-3.8-3.8
  • 'Five O'Clock Somewhere' 3.8-2-2
  • fluctuans 2-2-2
  • 'Follow Your Dream' 2.9-2.9-2.9
  • 'Forbidden Fruit' 4-4-4
  • 'Fortunei Albopicta' 2-2-2
  • 'Fourth of July' 2-2-2
  • 'Fragrant Blue' 2-2-2
  • 'Fragrant Bouquet' 2-2-2
  • 'Fragrant Gold' Aneuploid
  • 'Fran Godfrey' 3-3-3
  • 'Francee' 2-2-2
  • 'Frances Williams' 2-2-2
  • 'Friar Tuck' 3.5-3-3
  • 'Frosted Dimples' 4-4-4
  • 'Frosted June' 2-2-2
  • 'Frosted Mouse Ears' 2-2-2
  • 'Fujibotan' 2-2-2
  • 'Gaiety' 2-2-2
  • 'Gaijin' 2-2-2
  • 'Ginko Craig' 2-2-2
  • 'Gold Edger' 2-2-2
  • 'Gold Standard' 2-2-2
  • 'Golden Friendship' 2-2-2
  • 'Golden Memories' 2-2-2
  • 'Golden Needles' 4-4-4
  • 'Golden Tiara' 2-2-2
  • 'Goodness Gracious' 4-2-2
  • gracillima 2-2-2
  • gracillima 'Alba' 2-2-2
  • 'Grand Forks' 2-2-2
  • 'Grand Marquee'
    (1) 2-2-2
    (2) 4-2-2 (see footnote 4)
  • 'Grand Prize' 3.6-2-2
  • 'Grand Slam' 2-2-2
  • 'Grand Tiara'
    (1) 4-2-2
    (2) 4-4-4 (see footnote 4)
  • 'Great Escape' 4-2-2
  • 'Great Expectations' 2-2-2
  • 'Green Eyes' 2-2-2
  • 'Green Mouse Ears' 2-2-2
  • 'Guacamole' 2-2-2
  • 'Guardian Angel' 3-3-3
  • 'Gunther's Prize' 3-3-3
  • 'Gypsy Rose' 2-2-2
  • 'Hadspen Blue' 2-2-2
  • 'Haku Chu Han' 2-2-2
  • 'Hakuyo' 2-2-2
  • 'Halcyon' 2-2-2
  • 'Hands Up' 4-4-4
  • 'Happy Dayz' 2-2-2
  • 'Harry van de Laar' 2-2-2
  • 'Heat Wave' 4-2-2
  • 'Heaven Scent' 4-2-2
  • 'High Noon' 2-2-2
  • 'High Society' 1.9-1.9-2.1
  • 'Hime Tokudama' 2-2-2
  • 'Hirao Majesty' 2-2-2
  • 'Hirao Supreme' 2-2-2
  • 'Hiroa Tetra' 2-2-2
  • 'Holly's Dazzler' 2-2-2
  • 'Holly's Honey' 3-3-3
  • 'Honeybells' 2-2-2
  • 'Hudson Bay' 2-2-2
  • 'Hydon Sunset' 2-2-2
  • hypoleuca 2-2-2
  • 'Illicit Affair' 2-2-2
  • 'Inaho' 2-2-2
  • 'Independence' 4-4-4
  • 'Invincible' 2-2-2
  • 'Iris Frazier' 3-2-2
  • 'Iron Gate Delight' 2-2-2
  • 'Iron Gate Supreme' 2-2-2
  • 'Ivory Coast' 4-2-2
  • 'Iwa Soules' 2-2-2
  • 'Japanboy' 2-2-2
  • 'Japangirl' 2-2-2
  • 'Jaz' 2.4-2.4-2.4
  • 'Jim Cooper' Aneuploid
  • 'Jolly Green Dwarf' 4-4-4
  • jonesii 2-2-2
  • 'Joyce Trott' Aneuploid
  • 'June' 2-2-2
  • 'June Fever' 2-2-2
  • 'June Present' 2-2-2
  • 'Justine' 4-4-4
  • 'Kabitan' 2-2-2
  • 'Katsuragawa Beni' 2-2-2
  • 'Key Lime Pie' 2-2-2
  • 'Kifukurin Hyuga' 2-2-2
  • 'Kifukurin Ko Mame' 2-2-2
  • 'Kifukurin Ubatake' 2-2-2
  • kikutii 2-2-2
  • kikutii var. caput-avis 2-2-2
  • kikutii var. leuconata 2-2-2
  • kikutii var. polyneuron 2-2-2
  • kikutii var. tosana 2-2-2
  • kikutii var. yakusimensis 2-2-2
  • 'Kinba' 2-2-2
  • 'Kirishima' 2-2-2
  • kiyosumiensis 2-2-2
  • kiyosumiensis 'Busen' 2-2-2
  • kiyosumiensis 'Maiohgi' 2-2-2
  • kiyosumiensis 'Seiryu' 2-2-2
  • 'Kiwi Black Magic' 4-4-4
  • 'Korean Snow' 2-2-2
  • 'Koryu' 2-2-2
  • 'Lady Isobel Barnett' 3-3-3
  • 'Lakeside Black Satin' 4-4-4
  • 'Lakeside Coal Miner' 3-3-3
  • 'Lancifolia' 2-2-2
  • 'Last Dance' 3-3-4
  • 'Leather Sheen' Aneuploid
  • 'Lettuce Salad' 4-4-4
  • 'Liberty' 4-2-2
  • 'Liliput' 2-2-2
  • 'Lily Pad' 2-2-2
  • 'Linda Sue' 4-2-2
  • 'Lionheart' 2-2-2
  • 'Little Blue' 3-3-3 (see footnote 5)
  • 'Little Caesar' 2-2-2
  • 'Little Jay' 2-2-2
  • 'Little Jim' 2-2-2
  • 'Little Treasure' 4-2-2
  • 'Little Wonder' 2-2-2
  • 'Lodestar' 2.7-2.7-2.7
  • longipes 2-2-2
  • longipes 'Amagi Nishiki' 2-2-2
  • longipes 'Aurea' 2-2-2
  • longipes 'Gotemba Nishiki' 2-2-2
  • longipes f. hypoglauca 2-2-2
  • longipes 'Lancea' 2-2-2
  • longipes 'Maruba' 2-2-2
  • longipes 'Ogon' 2-2-2
  • longipes 'Tenryu Nishiki' 2-2-2
  • longissima 2-2-2
  • 'Love Pat' 2-2-2
  • 'Lovely Loretta' Aneuploid
  • 'Loyalist' 4-4-4
  • 'M. Owen Papuga' Aneuploid
  • 'Maekawa' 2-2-2
  • 'Magic Fire' 2-2-2
  • 'Majesty' 4-2-2
  • 'Mango Tango' 4-2.8-4
  • 'Marbled White' 2-2-2
  • 'Maruba Iwa' 2-2-2
  • 'Mary Marie Ann' 2-2-2
  • 'Masquerade' 2-2-2
  • 'Mata Hari' 2-2-2
  • 'Maui Buttercups' 2-2-2
  • 'Memories of Dorothy' 2-2-2
  • 'Metallic Sheen' Aneuploid
  • 'Middle Ridge' 4-2-2
  • 'Mighty Mouse' 2-2-2
  • 'Mikawa-no-yuki' 2-2-2
  • minor "Gosan" 2-2-2
  • 'Minuteman' 4-4-4
  • montana 2-2-2
  • montana 'Aureomarginata' 2-2-2
  • montana f. macrophylla 2-2-2
  • 'Moonstruck' 2-2-2
  • 'Morning Light' 4-4-4
  • 'Mount Kirishima' 2-2-2
  • 'Mourning Dove' 4-2-2
  • 'My Child Insook' 2-2-2
  • 'My Friend Bertha' 4-4-4
  • 'My Laura' 2-2-2
  • 'Nancy Gill'
    (1) 2.7-2.7-2.7
    (2) 2.8-2.8-2.8
  • 'Neat Splash' 2-2-2
  • 'Neutron Star' 4-4-4 (see footnote 6)
  • 'New Wave' Aneuploid
  • 'Night before Christmas' 4-2-2
  • 'Obsession' 2-2-2
  • 'Ogon Tsushima' 2-2-2
  • 'Oh My Heart' 2-2-2
  • 'Old Glory' 2-2-2
  • 'Olympic Sunrise' 4-2-2
  • 'On Stage' 2-2-2 (see footnote 7)
  • 'Opipara' 2-2-2
  • 'Orange Marmalade' 2-2-2
  • 'Orange Star' 4-4-4
  • 'Otome' 4-4-4
  • 'Otome-no-ka' 2-2-2
  • 'Outhouse Delight' 2-2-2
  • 'Pandora's Box' 2-2-2
  • 'Paradise Joyce' 2-2-2
  • 'Parhelion' 3-3-3
  • 'Pathfinder' 4-4-4
  • 'Patriot' 4-2-2 and 4-4-4 (see footnote 4)
  • 'PeeDee Elfin Bells' 3-3-3
  • 'Piedmont Gold' 2-2-2
  • 'Pitch Black' 4-4-4
  • plantaginea 2-2-2
  • 'Pocketful of Sunshine' 2-2-2
  • 'Pot of Gold' 2-2-2
  • 'Prairie Sum Shine' 3-3-3
  • 'Praying Hands' 2-2-2
  • 'Prestige and Promise' 3.8-3.8-2
  • 'Prince of Darkness' 4-4-4
  • pulchella 2-2-2
  • 'Purple Profusion' 2-2-2
  • pycnophylla 2-2-2
  • 'Radiant Edger' 4-4-4
  • 'Rainbow's End'
    (1) 2.5-2-2
    (2) 2-2-2.5
  • 'Rainforest Sunrise' 2-2-2
  • 'Raleigh Remembrance' 2.5-2.5-2.5
  • rectifolia 2-2-2
  • rectifolia 'Atropurpurea' 2-2-2
  • rectifolia 'Aurea' 2-2-2
  • rectifolia 'Sachalin' 2-2-2
  • rectifolia 'White Tachi' 2-2-2
  • 'Red Dragon' 2-2-2
  • 'Red Neck Heaven' 2-2-2
  • 'Red October' 2-2-2
  • 'Red Tubes' 2-2-2
  • 'Regal Rhubarb' 2-2-2
  • 'Remember Me' 2-2-2
  • 'Revolution' 4-4-4
  • 'Rhein' 2-2-2
  • 'Rhino Blue' 4-4-4
  • 'Rhino Gold' 2-2-2
  • 'Rhino Hide' 4-2-2
  • 'Rosedale Genevieve' Aneuploid
  • 'Rosemore' 2-2-2
  • 'Royal Standard' 2-2-2
  • 'Royal Super' 2-2-4
  • 'Royal Wedding' 2-2-2
  • 'Rumpelstilzchen' 2-2-2
  • rupifraga 2-2-2
  • 'Sagae' 2-2-2
  • 'Saikoku' 2-2-2
  • 'Saint Elmo's Fire' 2-2-2
  • 'Saishu Jima' 2-2-2
  • 'Second Wind' 4-4-4
  • 'Secret Ambition' 4-4-4
  • 'Setsurei' 2-2-2
  • 'Shade Master' 4-4-4
  • 'Shelleys' 2-2-2
  • 'Sherborne Swift' 2-2-2
  • shikokiana 2-2-2
  • 'Shining Tot' 2-2-2
  • 'Shiny Substance' 3-3-3
  • 'Sieboldiana 2-2-2
  • sieboldii 2-2-2
  • sieboldii 'Alba' 2-2-2
  • 'Silver Eagle' 2-2-2
  • 'Sitting Pretty' 2-2-2
  • 'Silver Threads & Golden Needles' 2-2-2
  • 'Sir Richard' 2-2-2
  • 'Sleeping Beauty' 2-2-2
  • 'Slick Willie' 2-2-2
  • 'Small Sum' Aneuploid
  • 'Snow Cap' 2-2-2
  • 'Snow Flakes' 2-2-2
  • 'Snow Mouse' 2-2-2
  • 'Snow White' 2-2-2
  • 'Something Good' 3-2.7-2.7
  • 'Sparkler' 3.2-3.2-3.2
  • 'Sparkling Burgundy' 2-2-2
  • 'Sparky' 2-2-2
  • 'Spilt Milk' 2-2-2
  • 'St. Paul' 4-2-2
  • 'Stand by Me' 2-2-2
  • 'Stargate' 2-2-2
  • 'Steady On' 2-2-2
  • 'Stiletto' 2-2-2
  • 'Stitch in Time' 4-2.5-3.6
  • 'Straka Gold' 2-2-2
  • 'Striptease' 2-2-2
  • 'Sum and Substance' 3-3-3
  • 'Sum and Subtle' 3-3-3
  • 'Sum Cup-o-Joe'
    (1) 3-3-3
    (2) 2.8-2.8-2.8
  • 'Sum It Up' 3-3-3
  • 'Sum of All' 3-2.7-2.5
  • 'Summer Breeze' 4-4-4
  • 'Summer Lovin' 4-4-4
  • 'Summer Music' 4-2-2
  • 'Swan Song' 2-2-2
  • 'Sweet Innocence' 4-2-2
  • 'Sweet Susan' 2-2-2
  • takahashii "Gosan" 2-2-2
  • 'Tardiflora' 2-2-2
  • 'Tardiflora Hybrida' 2-2-2
  • tardiva "Gosan" 2-2-2
  • 'Tea Time' 2-2-2
  • 'Tetra' 2-2-2
  • 'Theo's Blue' 2-2-2
  • tibae 2-2-2
  • 'Tiffney's Godzilla' 3-3-3
  • 'Tiny Tears' 2-2-2
  • 'Titanic' 3-3-3 (see footnote 7)
  • 'Tokudama' 2-2-2
  • 'Tokudama Aureonebulosa' 2-2-2
  • 'Tokudama Flavocircinalis' 2-2-2
  • 'Torchlight' 2-2-2
  • 'Tossed Salad' 2.3-2.3-2.3
  • 'Touch of Class'
    (1) 4-2-2
    (2) 4-4-4(see footnote 4)
  • 'Touch of Frost' 4-4-4
  • 'Trade Wind' 4-4-4
  • 'Treasure Island' 2-2-2
  • 'True North' 2.9-2.9-2.9
  • 'Tsugaru Komachi' 2-2-2
  • tsushimensis 2-2-2
  • 'Tucker Valentine' 3-3-3
  • 'Twilight' 4-4-4
  • 'Twilight Zone' 2-2-2
  • 'Ultraviolet Light' 2-2-2
  • 'Undulata Albomarginata' 2-2-2
  • 'Undulata Erromena' 2-2-2
  • 'Undulata Univittata' 2-2-2
  • 'Urajiro Hachijo' 2-2-2
  • 'Urajiro Hachyo' 2-2-2
  • 'Uzu-no-mai' 2-2-2
  • ventricosa 4-4-4
  • ventricosa 'Aureomaculata' 4-4-4
  • ventricosa 'Aureomarginata' 4-4-4
  • ventricosa 'Golden Spot' 4-4-4
  • ventricosa 'Schwarzer Ritter' 4-4-4
  • ventricosa 'Spring Gold' 4-4-4
  • 'Venucosa' 3-3-3
  • venusta 2-2-2
  • 'Vera Verde' 2-2-2
  • 'Victory' 2-2-2
  • 'Vim and Vigor' 2.8-2.8-2.8
  • 'Viridipes' 2-2-2
  • 'Volcano Island' 4-4-4
  • 'Vulcan' 4-2-2
  • 'Warwick Comet' 2-2-2
  • 'Warwick Essence' 4-4-4
  • 'Waukon Thin Ice' 2-2-2
  • 'Wheee!' 2-2-2
  • 'Whirlwind' 4-4-4
  • 'White Bikini' 2-2-2
  • 'White Tachi' 2-2-2
  • 'White Wall Tire' 2-2-2
  • 'William Lachman' 2-2-2
  • 'Windsor Blue' 2-2-2
  • 'Winter Snow' 3-3-3
  • 'Woop Woop' 4-4-4
  • 'Yakushima Mizu' 2-2-2
  • 'Yankee Blue' 2-2-2
  • 'Yea' 2-2-2
  • yingeri 2-2-2
  • 'Yoshinogawa' 2-2-2
  • 'Yu Lei' 2-2-2
  • 'Zebson' 3-3-3
  • 'Zero Sum' 2.9-2.9-2.9

Also see the "Polyploid Hostas" list.

1 According to Warren I. Pollock, "This and That: 2011," The Online Hosta Journal 42 Online, H. 'Bubba' is aneuploid, not diploid as previously stated by Steven C. Chamberlain, Warren I. Pollock, and Ben J. M. Zonneveld, "Ploidy and Pollen Viability in Hosta Cultivars: Zonneveld's Data Sets," The Hosta Journal 42, no. 1 (2011): 71-78.

2 According to Steven C. Chamberlain, "Ploidy and Pollen Viability in Hosta Cultivars: Zonneveld's Data Sets," The Hosta Journal 42, no. 1 (2011): 71-78, H. 'Cheatin Heart' is presented as a ploidy chimera (2-2-4) and a diploid (2-2-2). Upon re-examination of the flow cytometry data, it was determined that 'Cheatin Heart' is diploid (2-2-2).

3 Ben J. M. Zonneveld and Frank Van Iren, "Flow cytometric analysis of DNA content in Hosta reveals ploidy chimeras", Euphytica, 111, no. 2 (2000): 105-110 lists H. "Fire Works" as diploid. This hosta is not H. 'Fireworks' PP16,062 (Walters Gardens- 2001). Zonneveld and Van Iren, "H. 'Fireworks' and 'Sparkler': Unusual Aneuploid Sports from 'Loyalist'," The Hosta Journal 46, no. 2(2015): 72-76 states that H. 'Fireworks' (Walters Gardens- 2001) is aneuploid, 3.2-4-4.

4 According to Zonneveld and Frank Van Iren, "Flow cytometric analysis of DNA content," H. 'Patriot' is tetraploid. However, B. J. M. Zonneveld, "Nuclear DNA content of ploidy chimeras of Hosta Tratt. (Hostacae) demonstrate three apical layers in all organs, but not in the adventitious root," Plant Systematics and Evolution 269, no. 1 (2007): 29-38 states that H. 'Patriot' is partially tetraploid (4-2-2). Eventually, it was discovered that both partially tetraploid as well as fully tetraploid forms are in the trade.
  According to B. J. M. Zonneveld and Warren I. Pollock, "Flow cytometric analysis of somaclonal variation in lineages of Hosta sports detects polyploidy and aneuploidy chimeras," Plant Biology 14, no. 6 (2012): 972-979, "H. 'Patriot' exists as both partial tetraploid (4-2-2) and fully tetraploid (4-4-4) plants in the trade (Zonneveld 1999). Several other plants with different ploidy are in commerce under the same name. Examples are H. 'Grand Tiara', H. 'Grand Marquee' and H. 'Touch of Class', with both 4-2-2 and 4-4-4 in the trade. Seemingly, over the years, tissue culture propagation of 4-2-2 plants has given rise to 4-4-4 plants. As 4-2-2 plants have leaves comparable in thickness to 4-4-4 plants, this change might go unnoticed." Also see, Ben J. M. Zonneveld, "The DNA Content of H. 'Francee', 'Minuteman', and Two Different Varieties Called 'Patriot'," The Hosta Journal 30, no. 2 (1999): 28, 29.

5 According to Zonneveld and Frank Van Iren, "Flow cytometric analysis of DNA content," H. 'Little Blue' is incorrectly identified as a tetraploid (page 3) and an aneuploid (page 5). Zonneveld has since determined H. 'Little Blue' to be a triploid.

6 Hosta 'Neutron Star' was originally submitted as H. 'Otome' for flow cytometry measurement. See Steven C. Chamberlain, Warren I. Pollock, and Ben J. M. Zonneveld, "Ploidy and Pollen Viability in Hosta Cultivars: Zonneveld's Data Sets", The Hosta Journal 42, no. 1 (2011): 71-78. The hosta was determined to be 4-4-4 and the name was later changed to 'Neutron Star'.

7 According to Warren I. Pollock, "Reflections: 2008," The Hosta Journal 40, no. 1 (2009): 72-76, H. 'On Stage' is incorrectly presented as tetraploid. In Zonneveld and Van Iren, "Flow cytometric analysis of DNA content," H. 'On Stage' is confirmed as diploid.

8 Steven C. Chamberlain, Warren I. Pollock, and Ben J. M. Zonneveld, "Ploidy and Pollen Viability in Hosta Cultivars: Zonneveld's Data Sets," The Hosta Journal 42, no. 1 (2011): 71-78 incorrectly presents H. 'Titanic' as triploid. However, Zonneveld and Pollock, "Flow cytometric analysis of somaclonal variation," confirms H. 'Titanic' is aneuploid, 3-2.8-2.8.

Sources

Chamberlain, Steven C. "Zonneveld's Study of Cell Layers and Leaf Variegation in Hosta Cultivars: A Summary."
   The Hosta Journal, Volume 39, Number 2 (2008): 66-69.

Chamberlain, Steven C., Warren I. Pollock, and Ben J. M. Zonneveld. "Ploidy and Pollen Viability in Hosta Cultivars: Zonneveld's Data Sets."
   The Hosta Journal, Volume 42, Number 1 (2011): 71-78.

Mortko, Rob. "Hot Hostas, Chemically Induced Tets: A Sure Fire Method for Success?" The Online Hosta Journal, Volume 41 Online (2010).

Pollock, Warren I. Email to Donald Rawson regarding H. 'Majesty' and H. 'Victory'. February 21, 2015.

Pollock, Warren I. "Reflections: 2008." The Hosta Journal, Volume 40, Number 1 (2009): 72-76.

Pollock, Warren I. "This and That: 2011." The Online Hosta Journal, Volume 42 Online (2011).

Pollock, Warren I. "This and That: 2015; New Hosta Ploidy (L1-L2-L3) Measurements." The Online Hosta Journal, Volume 46 Online (2015).

Pollock, Warren I. "What's in a Hosta Name? Part LIII." The Hosta Journal, Volume 42, Number 2 (2011): 67-74.

Pollock, Warren I. "What's in a Hosta Name? Part LVII." The Hosta Journal, Volume 44, Number 2 (2013):8-18.

Pollock, Warren I and Ben J. M. Zonneveld. "More Information on Bob Solberg's 'Hybridizing thoughts 2015'."
    The Online Hosta Journal, Volume 46 Online (2015).

Solberg, Bob. "Do You Know a Tetraploid Hosta When You See One?" The Hosta Journal, Volume 42, Number 2 (2011): 33-36.

Solberg, Bob. Green Hill Farm. <http://www.hostahosta.com>.

Walek, Kevin. "Neutron Star". The Hosta Registry. 2011. <http://www.hostaregistrar.org>.

Zonneveld, Ben J. M. "Nuclear DNA content of ploidy chimeras of Hosta Tratt. (Hostaceae) demonstrate three apical layers in all organs, but not in the adventitious
   root." Plant Systematics and Evolution, Volume 269, Issue 1 (2007): 29-38.

Zonneveld, Ben J. M. "The DNA Content of H. 'Francee', 'Minuteman', and Two Different Varieties Called 'Patriot'."
   The Hosta Journal, Volume 30, Number 2 (1999): 28, 29.

Zonneveld, Ben J. M. and Frank Van Iren. "Flow cytometric analysis of DNA content in Hosta reveals ploidy chimeras."
   Euphytica, Volume 111, Issue 2 (2000): 105-110.

Zonneveld, Ben J. M. and Frank Van Iren. "Genome Size and Pollen Viability as Taxonomic Criteria: Application to the Genus Hosta."
   Plant Biology, Issue 3 (2001): 176-185.

Zonneveld, Ben J. M., Warren I. Pollock, Rob Mortko, and Steven C. Chamberlain. "H. 'Stitch in Time': A Multiple Mystery."
   The Hosta Journal, Volume 40, Number 1 (2009): 62-71.

Zonneveld, Ben J. M. and Warren I. Pollock. "Flow cytometric analysis of somaclonal variation in lineages of Hosta sports detects polyploidy and
    aneuploidy chimeras." Plant Biology, Volume 14, Issue 6 (2012): 972-979.

Zonneveld, Ben J. M. and Warren I. Pollock. "H. 'Fireworks' and 'Sparkler': Unusual Aneuploid Sports from 'Loyalist'."
   The Hosta Journal, Volume 46, Number 2 (2015): 72-76.

Zonneveld, Ben J. M. and Warren I. Pollock. "Sports and hybrids of triploid Hosta 'Sum and Substance' reveal chromosome losses and gains in all three apical
   layers." Plant Systematics and Evolution, Volume 298, Issue 6 (2012): 1037-1043.

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