Akagi Y, Akamatsu H, Otani H, Kodama M (2009a)
Horizontal chromosome transfer, a mechanism
for the evolution and differentiation of a plantpathogenic fungus. Eukaryot Cell 8:1732–1738.
https://doi.org/10.1128/EC.00135-09
Akagi Y, Taga M, Yamamoto M, Tsuge T, FukumasaNakai Y, Otani H, Kodama M (2009b) Chromosome constitution of hybrid strains constructed by
protoplast fusion between the tomato and strawberry pathotypes of Alternaria alternata. J Gen
Plant Pathol 75:101–109. https://doi.org/10.1007/
s10327-009-0149-1
Akamatsu H, Itoh Y, Kodama M, Otani H, Kohmoto K
(1997) AAL-toxin-deficient mutants of Alternaria
alternata tomato Pathotype by restriction enzymemediated integration. Phytopathology 87:967–972.
https://doi.org/10.1094/PHYTO.1997.87.9.967
Akamatsu H, Taga M, Kodama M, Johnson R, Otani H,
Kohmoto K (1999) Molecular karyotypes for Alternaria plant pathogens known to produce hostspecific toxins. Curr Genet 35:647–656
Akera T, Chma ´tal L, Trimm E, Yang K, Aonbangkhen
C, Chenoweth DM, Janke C, Schultz RM, Lampson
MA (2017) Spindle asymmetry drives nonMendelian chromosome segregation. Science
358:668–672.
https://doi.org/10.1126/science.
aan0092
Baayen RP, O’Donnell K, Bonants PJ, Cigelnik E, Kroon
LP, Roebroeck EJ, Waalwijk C (2000) Gene genealogies and AFLP analyses in the Fusarium oxysporum complex identify monophyletic and
nonmonophyletic Formae Speciales causing wilt
and rot disease. Phytopathology 90:891–900.
https://doi.org/10.1094/PHYTO.2000.90.8.891
Balesdent M-H, Fudal I, Ollivier B, Bally P, Grandaubert
J, Eber F, Che `vre A-M, Leflon M, Rouxel T (2013)
The dispensable chromosome of Leptosphaeria
maculans shelters an effector gene conferring
avirulence towards Brassica rapa. New Phytol
198:887–898. https://doi.org/10.1111/nph.12178
Banaei-Moghaddam AM, Martis MM, Macas J, Gundlach H, Himmelbach A, Altschmied L, Mayer
KFX, Houben A (2015) Genes on B chromosomes:
old questions revisited with new tools. Biochim
Biophys Acta 1849:64–70. https://doi.org/10.1016/
j.bbagrm.2014.11.007
Banaei-Moghaddam AM, Meier K, Karimi-Ashtiyani R,
Houben A (2013) Formation and expression of
pseudogenes on the B chromosome of rye. Plant
Cell
25:2536–2544.
https://doi.org/10.1105/
tpc.113.111856
Banaei-Moghaddam AM, Schubert V, Kumke K, Weib
O, Klemme S, Nagaki K, Macas J, Gonza ´lez-Sa ´nchez M, Heredia V, Go ´mez-Revilla D, Gonza ´lezGarcı ´a M, Vega JM, Puertas MJ, Houben A (2012)
Nondisjunction in favor of a chromosome: the
mechanism of rye B chromosome drive during
pollen mitosis. Plant Cell 24:4124–4134. https://
doi.org/10.1105/tpc.112.105270
Bauerly E, Hughes SE, Vietti DR, Miller DE, McDowell
W, Hawley RS (2014) Discovery of supernumerary
B chromosomes in Drosophila melanogaster.
Genetics 196:1007–1016. https://doi.org/10.1534/
genetics.113.160556
Bertazzoni S, Williams A, Jones DAB, Syme RA, Tan KC, Hane JK (2018) Accessories make the outfit:
accessory chromosomes and other dispensable
DNA regions in plant-pathogenic fungi. Mol
Plant-Microbe Interact 8:779–788. https://doi.org/
10.1094/MPMI-06-17-0135-FI
Bougourd SM, Jones RN (1997) B chromosomes: a
physiological enigma. New Phytol 137:43–54.
https://doi.org/10.1046/j.1469-8137.1997.00823.x
Camacho JPM, Schmid M, Cabrero J (2011) B chromosomes and sex in animals. Sex Dev 5:155–166.
https://doi.org/10.1159/000324930
Carchilan M, Delgado M, Ribeiro T, Costa-Nunes P,
Caperta A, Morais-Cecı ´lio L, Jones RN, Viegas
W, Houben A (2007) Transcriptionally active
heterochromatin in rye B chromosomes. Plant
Cell
19:1738–1749.
https://doi.org/10.1105/
tpc.106.046946
Cheng Y-M (2010) Evolution of the heterochromatic
regions on maize B long arm based on the
sequence structure of CL-repeat variants. Chromosom Res 18:605–619. https://doi.org/10.1007/
s10577-010-9136-9
Chuma I, Isobe C, Hotta Y, Ibaragi K, Futamata N,
Kusaba M, Yoshida K, Terauchi R, Fujita Y, Nakayashiki H, Valent B, Tosa Y (2011) Multiple translocation of the AVR-Pita effector gene among
chromosomes of the rice blast fungus Magnaporthe
oryzae and related species. PLoS Pathog 7:e1002147.
https://doi.org/10.1371/journal.ppat.1002147
Chuma I, Tosa Y, Taga M, Nakayashiki H, Mayama S
(2003) Meiotic behavior of a supernumerary chromosome in Magnaporthe oryzae. Curr Genet
43:191–198. https://doi.org/10.1007/s00294-0030390-7
Coleman JJ, Rounsley SD, Rodriguez-Carres M, Kuo A,
Wasmann CC, Grimwood J, Schmutz J, Taga M,
White GJ, Zhou S, Schwartz DC, Freitag M, Ma L-J,
Danchin EGJ, Henrissat B, Coutinho PM, Nelson
DR, Straney D, Napoli CA, Barker BM, Gribskov
M, Rep M, Kroken S, Molna ´r I, Rensing C, Kennell
JC, Zamora J, Farman ML, Selker EU, Salamov A,
Shapiro H, Pangilinan J, Lindquist E, Lamers C,
Grigoriev IV, Geiser DM, Covert SF, Temporini E,
Vanetten HD (2009) The genome of Nectria
haematococca: contribution of supernumerary
chromosomes to gene expansion. PLoS Genet
5:e1000618.
https://doi.org/10.1371/journal.
pgen.1000618
Coleman JJ, Wasmann CC, Usami T, White GJ, Temporini ED, McCluskey K, Vanetten HD (2011)
Characterization of the gene encoding pisatin
demethylase (FoPDA1) in Fusarium oxysporum.
2 Origin, Function, and Transmission of Accessory Chromosomes
41
Horizontal chromosome transfer, a mechanism
for the evolution and differentiation of a plantpathogenic fungus. Eukaryot Cell 8:1732–1738.
https://doi.org/10.1128/EC.00135-09
Akagi Y, Taga M, Yamamoto M, Tsuge T, FukumasaNakai Y, Otani H, Kodama M (2009b) Chromosome constitution of hybrid strains constructed by
protoplast fusion between the tomato and strawberry pathotypes of Alternaria alternata. J Gen
Plant Pathol 75:101–109. https://doi.org/10.1007/
s10327-009-0149-1
Akamatsu H, Itoh Y, Kodama M, Otani H, Kohmoto K
(1997) AAL-toxin-deficient mutants of Alternaria
alternata tomato Pathotype by restriction enzymemediated integration. Phytopathology 87:967–972.
https://doi.org/10.1094/PHYTO.1997.87.9.967
Akamatsu H, Taga M, Kodama M, Johnson R, Otani H,
Kohmoto K (1999) Molecular karyotypes for Alternaria plant pathogens known to produce hostspecific toxins. Curr Genet 35:647–656
Akera T, Chma ´tal L, Trimm E, Yang K, Aonbangkhen
C, Chenoweth DM, Janke C, Schultz RM, Lampson
MA (2017) Spindle asymmetry drives nonMendelian chromosome segregation. Science
358:668–672.
https://doi.org/10.1126/science.
aan0092
Baayen RP, O’Donnell K, Bonants PJ, Cigelnik E, Kroon
LP, Roebroeck EJ, Waalwijk C (2000) Gene genealogies and AFLP analyses in the Fusarium oxysporum complex identify monophyletic and
nonmonophyletic Formae Speciales causing wilt
and rot disease. Phytopathology 90:891–900.
https://doi.org/10.1094/PHYTO.2000.90.8.891
Balesdent M-H, Fudal I, Ollivier B, Bally P, Grandaubert
J, Eber F, Che `vre A-M, Leflon M, Rouxel T (2013)
The dispensable chromosome of Leptosphaeria
maculans shelters an effector gene conferring
avirulence towards Brassica rapa. New Phytol
198:887–898. https://doi.org/10.1111/nph.12178
Banaei-Moghaddam AM, Martis MM, Macas J, Gundlach H, Himmelbach A, Altschmied L, Mayer
KFX, Houben A (2015) Genes on B chromosomes:
old questions revisited with new tools. Biochim
Biophys Acta 1849:64–70. https://doi.org/10.1016/
j.bbagrm.2014.11.007
Banaei-Moghaddam AM, Meier K, Karimi-Ashtiyani R,
Houben A (2013) Formation and expression of
pseudogenes on the B chromosome of rye. Plant
Cell
25:2536–2544.
https://doi.org/10.1105/
tpc.113.111856
Banaei-Moghaddam AM, Schubert V, Kumke K, Weib
O, Klemme S, Nagaki K, Macas J, Gonza ´lez-Sa ´nchez M, Heredia V, Go ´mez-Revilla D, Gonza ´lezGarcı ´a M, Vega JM, Puertas MJ, Houben A (2012)
Nondisjunction in favor of a chromosome: the
mechanism of rye B chromosome drive during
pollen mitosis. Plant Cell 24:4124–4134. https://
doi.org/10.1105/tpc.112.105270
Bauerly E, Hughes SE, Vietti DR, Miller DE, McDowell
W, Hawley RS (2014) Discovery of supernumerary
B chromosomes in Drosophila melanogaster.
Genetics 196:1007–1016. https://doi.org/10.1534/
genetics.113.160556
Bertazzoni S, Williams A, Jones DAB, Syme RA, Tan KC, Hane JK (2018) Accessories make the outfit:
accessory chromosomes and other dispensable
DNA regions in plant-pathogenic fungi. Mol
Plant-Microbe Interact 8:779–788. https://doi.org/
10.1094/MPMI-06-17-0135-FI
Bougourd SM, Jones RN (1997) B chromosomes: a
physiological enigma. New Phytol 137:43–54.
https://doi.org/10.1046/j.1469-8137.1997.00823.x
Camacho JPM, Schmid M, Cabrero J (2011) B chromosomes and sex in animals. Sex Dev 5:155–166.
https://doi.org/10.1159/000324930
Carchilan M, Delgado M, Ribeiro T, Costa-Nunes P,
Caperta A, Morais-Cecı ´lio L, Jones RN, Viegas
W, Houben A (2007) Transcriptionally active
heterochromatin in rye B chromosomes. Plant
Cell
19:1738–1749.
https://doi.org/10.1105/
tpc.106.046946
Cheng Y-M (2010) Evolution of the heterochromatic
regions on maize B long arm based on the
sequence structure of CL-repeat variants. Chromosom Res 18:605–619. https://doi.org/10.1007/
s10577-010-9136-9
Chuma I, Isobe C, Hotta Y, Ibaragi K, Futamata N,
Kusaba M, Yoshida K, Terauchi R, Fujita Y, Nakayashiki H, Valent B, Tosa Y (2011) Multiple translocation of the AVR-Pita effector gene among
chromosomes of the rice blast fungus Magnaporthe
oryzae and related species. PLoS Pathog 7:e1002147.
https://doi.org/10.1371/journal.ppat.1002147
Chuma I, Tosa Y, Taga M, Nakayashiki H, Mayama S
(2003) Meiotic behavior of a supernumerary chromosome in Magnaporthe oryzae. Curr Genet
43:191–198. https://doi.org/10.1007/s00294-0030390-7
Coleman JJ, Rounsley SD, Rodriguez-Carres M, Kuo A,
Wasmann CC, Grimwood J, Schmutz J, Taga M,
White GJ, Zhou S, Schwartz DC, Freitag M, Ma L-J,
Danchin EGJ, Henrissat B, Coutinho PM, Nelson
DR, Straney D, Napoli CA, Barker BM, Gribskov
M, Rep M, Kroken S, Molna ´r I, Rensing C, Kennell
JC, Zamora J, Farman ML, Selker EU, Salamov A,
Shapiro H, Pangilinan J, Lindquist E, Lamers C,
Grigoriev IV, Geiser DM, Covert SF, Temporini E,
Vanetten HD (2009) The genome of Nectria
haematococca: contribution of supernumerary
chromosomes to gene expansion. PLoS Genet
5:e1000618.
https://doi.org/10.1371/journal.
pgen.1000618
Coleman JJ, Wasmann CC, Usami T, White GJ, Temporini ED, McCluskey K, Vanetten HD (2011)
Characterization of the gene encoding pisatin
demethylase (FoPDA1) in Fusarium oxysporum.
2 Origin, Function, and Transmission of Accessory Chromosomes
41