FGSC #2581

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Reporting Genes: his-2 ad-3A ad-3B nic-2;inl;ad-2+al-2;cot-1;pan-2

Species: crassa

Allele: (C94 Y112M13 35203 43002 JH319 Y175M256)+(Y112M38 C102(t) Y387-15.7)

Alternate Strain Number: Heterokaryon

Depositor: FJD

Linkage Group: (IR;IIIR;VR) + (IR;IVR;VIR)

Mating Type: A

Species Number: 10

Deposit Sheet ↗

Genes

Locus	Cultural Requirements	Link Group	Type
al-2	IR. Right of os-5 (<1%) and T(STL76). Left of arg-6(1%) and al-1 (797, 802, 808, 816, 818). Included in duplications from Tp(T54M94), confirming location left of arg-6(808). (482) Carotenoids absent or abnormal, but steroids produced (398). Blocked in microsomal fraction and defective in phytoene synthetase (445), a particulate enzyme (445 and references cited therein) (Fig. 9). Tracer experiments indicate a lesion between prephytoene pyrophosphate and phytoene (572). Alleles include those resulting in white and pale rose-white, e.g., 15300 and Y254MI65 (1042), and purple, e.g., MN58a (154). For complementation, see references 500 and 1041. Fine-structure mapping (500, 1042) needs reevaluation because of new information on the location of the arg-6 marker (797).	IR	B
pan-2	VIR. Right of rib-1 (<1 to 3%). Left of del (6%) and trp-2(11%) (140, 141, 143, 818, PB). Unable to convert ketovaline to ketopantoic acid (138, 140, 141). Used in major studies of intralocus recombination and complementation (140-143). pan-2ascospores remain white or pale if the crossing medium is not supplemented, even when the protoperithecial parent is pan-2+. Asci in which gene conversion has occurred at pan-2 can thus be recognized and isolated (1072, 1073); photographs (1072). For good recovery of pan-2progeny, crossing media should be supplemented with pantothenic acid (10 µg/ml) even when the protoperithecial parent is pan+. Called group B.	VIR	B
nic-2	IR. Between ad-3B (4%) and ace-7 (4 to 7%) (271, 578). (482) Grows on nicotinic acid, nicotinamide, or high concentrations of quinolinic acid (97, 1168). Cannot use kynurenine, hydroxykynurenine, or hydroxyanthranilic acid (96, 1168). Accumulates 3-hydroxyanthranilic acid (96) (Fig. 18). Aging cultures accumulate red-brown pigment in the medium. Used to study intralocus recombination (908). Translocations T(4540) and T(S1325)are inseparable from nic-2 (808, 908, 911).	IR	B
inl	VR. Between pho-3 (3 to 4%) and pab-1 (1 to 10%). Right of al-3 (362, 397, 1036). (482)Requires inositol (65). Lacks D-myoinositol-1-phosphatase (1142). Lack of glucocycloaldolase found by Pina and Tatum (826) is attributed by Williams (1142) to drastic repression of glucocycloaldolase by the concentration of inositol used for growth. Growth is colonial on low levels of inositol (367). Tends to extrude dark pigment into the medium when grown on suboptimal inositol. Composition of phospholipids and cell walls is abnormal on limiting inositol (367, 439, 440, 501). Inhibited by hexachlorocyclohexane (366, 457, 931). Conidia are subject to death by unbalanced growth on minimal medium (1028, 1033), a property exploited for mutant enrichment ("inositol-less death") (606, 647) because double mutants are at a selective advantage. Heat-sensitive allele 83201 is especially useful for mutant enrichment (832, 1043). Used in the first experiments reporting transformation of Neurospora by N. crassaDNA (677, 679) and reported to be efficient as a recipient in absence of inositol (1162). Used to study glucose (917) and sulfate (641) transport systems. Used extensively for studying induced reversion (392). Used for studying the mechanism of inositol-less death (647, 702), mutagenicity of ferrous ions, and regulation of mitochondrial membrane fluidity; for a review, see reference 702. Spontaneous reversion rates (386). Allele-specific partial suppressor (390). Allele 46802 is nonrevertable and inseparable from translocation 46802 (386, 808). Strains carrying heat-sensitive allele 83201 show slow semicolonial growth in liquid minimal medium at 25°C (641), but look normal on slants (D.D. Perkins, unpublished data). Strains carrying allele 89601 contain cross-reacting material (1183). Mutant gene exo-1 is present in the inl(89601) a stock FGSC 498 and may, therefore, be present in stocks of mutants derived by inositol-less death. (See references 194, 325, and 1027). Called inos.	VR	B
cot-1	IVR. Between pan-1 (2%) and his-4 (1 to 6%) (692, 812, 816). Extremely colonial at 34°C, but completely normal growth, morphology, and fertility at 25°C and below. Linear growth is maximum at 24°C (374). Becomes colonial at 32°C; colonies from ascospores or conidia are viable and continue to grow slowly with dense branching, but do not conidiate. They quickly resume normal growth when shifted to a permissive temperature (692, 1068). Recessive in duplications (808); apparent dominance in heterokaryons (374) may have resulted from a shift in nuclear ratios. Used in studies of septation and branching (202), growth-inhibiting mucopolysaccharide (878, 879), and sulfate transport (641). Cell wall analysis (374). Growth is stimulated by lysine or arginine (0.1 mM) on glucose media at high temperatures (615). Because of high viability and tightly restricted growth at restrictive temperatures and normality at 25°C, cot-1 mutants have valuable technical applications. For example, crosses homozygous for cot-1 have been used in combination with sorbose for experiments with rec genes, where high-density ascospore platings are required for precise quantitative analysis of intralocus recombination (e.g., references 165, 997, and 1070). In another application, when shifted up after initial growth at the permissive low temperature, cot-1hyphae assume a "bottle brush" appearance with small side branches (692). This has been used to select uvs mutants by subsurface survival on UV-irradiated plates containing p-aminobenzoic acid (938; D.E.A. Catcheside, personal communication). cot-1 conidia or ascospores from cot-1 x cot-1crosses are used for replication in a protocol involving transfer by filter paper (615). For suppressors of cot-1, see gul.	IVR	B
ad-3B	IR. Between ad-3A (0.1 to 0.7%) and nic-2 (3%) (271). (482) Uses adenine or hypoxanthine (682). Blocked in interconversion of AIR to CAIR (348) (Fig. 8). Produces purple pigment, permitting direct visual selection (276, 682). Pigment is secreted with low concentrations of adenine (e.g., 0.1 mM), not with high concentrations (2 mM) (276, 682, 785). Pigment production used to assess effect of histidine and tryptophan on purine nucleotide synthesis (786). Reduced interallelic fertility (264, 407). Complementation maps (268, 274). Relation of mutagens to complementation patterns (269). Mutants with non-polarized complementation patterns on the right side of the complementation map grow on minimal medium if supplied with CO2; other mutants do not respond to CO2, (270). Used extensively for mutagenesis (see ad-3A). Rearrangement T(I- >III)Y112M4i ad-3B, which has a breakpoint inseparable from ad-3B, was the first insertional translocation to be reported for fungi (266). Allele 7-017-0137 shows "fixed instability," mutating to an unstable prototrophic allele (41). Alleles 2-17-126, 12-21-28, and numerous others are supersuppressible (408, 749, 955). Called complementation group B.	IR	B
ad-3A	IR. Between his-3 (1 to 2%) and ad-3B (0.1 to 0.7%) (271). Right of ure-4 (78). (482) Requires adenine or hypoxanthine (682). Blocked in interconversion of CAIR plus aspartate to SAICAR (348) (Fig. 8). Produces purple pigment, permitting direct visual selection (276, 682); see the ad-3B entry. Reduced interallelic fertility (407). No interallelic complementation (267; F.J. de Serres, personal communication). ad-3A and ad-3B are two genetically and functionally distinct loci separated by a short but functionally complex region of unknown but essential function (271, 407). They have been used intensively for quantitative genetic and molecular studies of mutation (for a review, see reference 35). Either forward mutation (e.g., reference 277) or reverse mutation (e.g., reference 772) can be measured precisely; the former is detected visually by purple pigment. Purple pigment has also been used to assess the effect of histidine and tryptophan on purine nucleotide synthesis (786). Alleles N23 and N24 have been used as mutagen testers. N23 reverts with agents that cause base pair substitutions; N24 reverts with agents that cause frameshifts (772). SK(ad-3A) is at or near ad-3A and may be a cryptic ad-3A allele. Does not require adenine. In SK(ad-3A)x ad-3Acrosses, the ad-3A progeny die; possibly SK(ad-3A) mutants fail to make enough adenine to support their growth (251). Translocations Y155M64 ad-3A (272; PB) and Y112M15 ad-3A(413) each have one breakpoint that is inseparable from ad-3A. Called complementation group A (264). "A" in the locus symbol does not refer to mating type.	IR	B
ad-2	IIIR. Between thi-2 (1%) and trp-1 (1 to 7%) (11, 219). (482) Requires adenine or hypoxanthine (682). Controls conversion of phosphoribosylformylglycineamidine to AIR (120) (Fig. 8). Strains carrying allele 70004(t) are heat sensitive (34°C versus 25 C) (682) and osmotic remediable (636). Called complementation group H.	IIIR	B
his-2	IR. Right of T(AR190) and un-2 (<1%). Left of the T(AR173) right breakpoint and of nuc-1 (<1%) (172, 670, 808). (434)Requires histidine (434). Affects adenosine 5'-triphosphate phosphoribosylpyrophosphate pyrophosphorylase (16) (Fig. 14). Intralocus complementation (162). Recombination between his-2 alleles is controlled by rec-3(173); it is not affected by rec-1 (172). Initial his-2 allele called C94.	IR	B