White haze
Figure 1. Apples of the cultivar Elstar showing signs of white haze.

Fig. 3. Scanning electron microscopy of apple with white haze, showing distinct
growth of filamentous hyphae of the anamorphic basidiomycetous genus
Tilletiopsis and Tilletiopsis-anamorphs of Entyloma. Insert shows sterigma of on
which ballistoconidia are being formed (arrow) Bar indicates 10µm.





RAPD-patterns
RAPD-patterns of ballistoconidium-forming fungi (Tilletiopsis and Entyloma spp.)
isolated from White haze showing the presence of genetic polymorphisms. A,
(ATG)5; B, (GTG)5; C, ERIC 1E; D, ERIC 2; E, OPA5.

Tabel
Table 1.
Origin of Tilletiopsis and
Entyloma isolates obtained from patches with
white haze on apples from The Netherlands (WH = White haze; R =
Russeted, PD = Plant Protection Service,
Wageningen, The Netherlands)
|
Name
|
CBS number
|
Origin
|
Russetted
/WH
|
Apple Cultivar
|
Year
|
|
Entyloma aff. matricariae
|
111599
|
Orchard Verkoogen
|
R
|
Elstar
|
1998
|
|
Entyloma
aff. matricariae
|
111600
|
Orchard Krabbendijk
|
R
|
Elstar
|
1998
|
|
Entyloma
aff. matricariae
|
111602
|
Orchard Dronten
|
R
|
Elstar
|
1998
|
|
Entyloma
aff. matricariae
|
111597
|
Orchard
Zeewolde
|
R
|
Elstar
|
1998
|
|
Entyloma
aff. matricariae
|
111598
|
Orchard Biddinghuizen
|
R
|
Elstar
|
1998
|
|
Entyloma
aff. matricariae
|
111601
|
Orchard Zeewolde
|
R
|
Elstar
|
1998
|
|
Entyloma
aff. matricariae
|
111596
|
Storage
|
WH
|
Elstar
|
1998
|
|
Entyloma spec. A
|
111603
|
Storage
|
WH
|
Elstar
|
1998
|
|
Entyloma
spec. A
|
111604
|
PD
|
WH
|
Elstar
|
1994
|
|
Entyloma
spec. A
|
111627
|
PD
|
WH
|
Elstar
|
1994
|
|
Entyloma spec. B
|
111607
|
Orchard Oostkapelle
|
?
|
Elstar
|
1998
|
|
Entyloma
spec. C
|
111594
|
PD
|
WH
|
Elstar
|
1994
|
|
Entyloma
spec. C
|
111595
|
Orchard Biddinghuizen
|
R
|
Elstar
|
1998
|
|
Entyloma
spec. C
|
111593
|
Storage
|
WH
|
Elstar
|
1998
|
|
Entyloma spec. D
|
111605
|
Orchard Wognum
|
?
|
Elstar
|
1998
|
|
Entyloma
spec. D
|
111606
|
Orchard Wognum II
|
?
|
?
|
1998
|
|
Tilletiopsis minor
|
111629
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis minor
|
111630
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis minor
|
111631
|
Orchard Wognum II
|
?
|
?
|
1998
|
|
|
111622
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis
pallescens
|
111623
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis
pallescens
|
111626
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis
pallescens
|
111624
|
Orchard
Wognum II
|
?
|
?
|
1998
|
|
Tilletiopsis
spec. A
|
111628
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis
spec. A
|
111625
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis
spec. B
|
111616
|
Storage
|
WH
|
Elstar
|
1998
|
|
Tilletiopsis
spec. B
|
111617
|
Storage
|
WH
|
Elstar
|
1998
|
|
Tilletiopsis
spec. B
|
111613
|
Storage
|
R
|
Elstar
|
1998
|
|
Tilletiopsis
spec. B
|
111615
|
Storage
|
R
|
Elstar
|
1998
|
|
Tilletiopsis
spec. B
|
111608
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis
spec. B
|
-
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis spec. B
|
111610
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis spec. B
|
111611
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis spec. B
|
111741
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis spec. B
|
111620
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis spec. B
|
-
|
PD
|
WH
|
Elstra
|
1994
|
|
Tilletiopsis spec. B
|
111612
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis spec. B
|
111609
|
PD
|
WH
|
Elstar
|
1994
|
|
Tilletiopsis spec. B
|
111619
|
Orchard
Wognum II
|
?
|
?
|
1998
|
|
Tilletiopsis spec. B
|
111618
|
Orchard
Wognum II
|
?
|
?
|
1998
|
|
Tilletiopsis spec. B
|
111614
|
Orchard Wognum II
|
?
|
?
|
1998
|
|
Tilletiopsis spec. B
|
111621
|
Orchard
Wognum II
|
?
|
?
|
1998 |
Cutinase, pectinase, protease and cellulase activity of Tilletiopsis and
Entyloma isolates from White haze (E. = Entyloma, T. =
Tilletiopsis, Tw = Tween, + = positive; ++ strongly positive; +++ = very
strongly positive; w= weak reaction; - = no activity; ? = not tested).
|
Name
|
Isolate
|
Tw 40
|
Tw 60
|
Tw 80
|
Cutinase
|
Pectinase
|
Protease
|
Cellulase
|
|
|
T. minor Nyland
|
CBS111629
|
+
|
+
|
+
|
+
|
+
|
++
|
W
|
|
|
T. minor Nyland
|
CBS111630
|
+
|
+
|
+
|
+
|
+
|
+
|
W
|
|
|
T. minor Nyland
|
CBS111631
|
+
|
+
|
+
|
+
|
+
|
++
|
W
|
|
|
T. pallescens Gokhale
|
CBS111622
|
+
|
+
|
+
|
+
|
++
|
++
|
-
|
|
|
T. pallescens Gokhale
|
CBS111623
|
+
|
+
|
+
|
+
|
+
|
+
|
-
|
|
|
T. pallescens Gokhale
|
CBS111624
|
+
|
+
|
+
|
+
|
++
|
++
|
W
|
|
|
T. pallescens Gokhale
|
CBS111626
|
+
|
+
|
+
|
+
|
++
|
++
|
W
|
|
|
T. species A
|
CBS111628
|
+
|
+
|
+
|
+
|
++
|
++
|
-
|
|
|
T. species A
|
CBS111625
|
+
|
+
|
+
|
+
|
++
|
++
|
-
|
|
|
T. species B
|
CBS111616
|
+
|
+
|
+
|
+
|
+++
|
++
|
-
|
|
|
T. species B
|
CBS111617
|
+
|
+
|
+
|
+
|
+++
|
++
|
W
|
|
|
T. species B
|
CBS111613
|
+
|
+
|
+
|
+
|
+
|
+
|
-
|
|
|
T. species B
|
CBS111615
|
+
|
+
|
+
|
+
|
+++
|
++
|
W
|
|
|
T. species B
|
CBS111608
|
+
|
+
|
+
|
+
|
+
|
+
|
-
|
|
|
T. species B
|
CBS111610
|
+
|
+
|
+
|
+
|
+++
|
++
|
-
|
|
|
T. species B
|
CBS111611
|
+
|
+
|
+
|
+
|
+++
|
++
|
-
|
|
|
T. species B
|
CBS111741
|
+
|
+
|
+
|
+
|
+++
|
++
|
W
|
|
|
T. species B
|
CBS111620
|
+
|
+
|
+
|
+
|
+
|
+
|
W
|
|
|
T. species B
|
CBS111612
|
+
|
+
|
+
|
+
|
+++
|
++
|
-
|
|
|
T. species B
|
CBS111609
|
+
|
+
|
+
|
+
|
+++
|
++
|
-
|
|
|
T. species B
|
CBS111619
|
+
|
+
|
+
|
+
|
+
|
++
|
+
|
|
|
T. species B
|
CBS111618
|
+
|
+
|
+
|
+
|
+++
|
++
|
-
|
|
|
T. species B
|
CBS111614
|
+
|
+
|
+
|
+
|
+
|
++
|
W
|
|
|
T. species B
|
CBS111621
|
+
|
+
|
+
|
+
|
+++
|
++
|
+
|
|
|
T. species C
|
CBS111607
|
+
|
+
|
+
|
+
|
+
|
++
|
-
|
|
|
T. species D
|
CBS111594
|
+
|
+
|
+
|
+
|
++
|
++
|
+
|
|
|
T. species D
|
CBS111595
|
+
|
+
|
+
|
+
|
+
|
+
|
w
|
|
|
T. species
E
|
CBS111603
|
+
|
+
|
+
|
+
|
+
|
++
|
+
|
|
|
T. species E
|
CBS111604
|
+
|
+
|
+
|
?
|
+
|
+
|
+
|
|
|
T. species F
|
CBS111599
|
+
|
+
|
+
|
+
|
++
|
++
|
-
|
|
|
T. species F
|
CBS111600
|
+
|
+
|
+
|
?
|
+
|
+
|
-
|
|
|
T. species F
|
CBS111602
|
+
|
+
|
+
|
+
|
+
|
++
|
W
|
|
|
T. species F
|
CBS111597
|
+
|
+
|
+
|
+
|
+
|
++
|
-
|
|
|
T. species F
|
CBS111598
|
+
|
+
|
+
|
?
|
+
|
+
|
-
|
|
|
T. species F
|
CBS111601
|
+
|
+
|
+
|
+
|
++
|
++
|
W
|
|
|
T. species F
|
CBS111596
|
+
|
+
|
+
|
+
|
+
|
+
|
W
|
|
|
T. species F
|
CBS111627
|
++
|
++
|
++
|
+
|
+
|
+
|
-
|
|
|
T. species
G
|
CBS111605
|
+
|
+
|
+
|
+
|
++
|
++
|
-
|
|
|
T. species
G
|
CBS111606
|
+
|
+
|
+
|
+
|
+
|
++
|
W
|
|
Genotypic variation within white haze fungi and reference species as analyzed by
RAPD and AFLP analysis. RAPD patterns are indicated with a letter, if they
differ in only a single band this is indicated by adding an additional symbol,
AFLP clusters are indicated with a figure, subclusters are indicated with an
additional ‘. Legends: PD = Plant protection service, Wageningen, The
Netherlands; E. = Entyloma; T. = Tilletiopsis; Ti. = Tilletia;
aff. = affinity to; spec. = species).
|
Name.
|
Isolate
|
AFLP cluster
|
RAPD GTG5
|
RAPD
ATG5
|
RAPD
Eric-2
|
RAPD
OPA-05
|
RAPD
Eric-1E
|
Overall
RAPD pattern
|
|
T. albescens
|
CBS 608.83
|
?
|
D
|
D
|
D
|
D
|
D
|
D
|
|
Ti. ayresii
|
CBS 188.97
|
?
|
E
|
?
|
E
|
E
|
?
|
E
|
|
T. flava
|
CBS 401.84
|
?
|
H
|
H
|
H
|
H
|
H
|
H
|
|
T. fulvescens
|
CBS 607.83
|
?
|
F
|
F
|
F
|
F
|
?
|
F
|
|
T. minor
|
CBS 543.50
|
?
|
A
|
?
|
A
|
A
|
A
|
A
|
|
T. minor Nyland
|
CBS111629
|
8
|
S
|
S
|
S
|
S
|
S
|
S
|
|
T. minor Nyland
|
CBS111630
|
8
|
S
|
S
|
S^^
|
S
|
S
|
S^^
|
|
T. minor Nyland
|
CBS111631
|
8
|
S*
|
S*
|
S*
|
S*
|
S*
|
S*
|
|
T. pallescens
|
CBS 606.83
|
?
|
G
|
G
|
G
|
G
|
G
|
G
|
|
T. pallescens
|
CBS
111623
|
4
|
O*
|
O*
|
O*
|
O*
|
O*
|
O*
|
|
T. pallescens Gokhale
|
CBS111626
|
4
|
O^^+
|
O*
|
O*
|
O*
|
O*
|
O
|
|
T. pallescens Gokhale
|
CBS111624
|
6
|
O^^
|
O^^
|
O^^
|
O^^
|
O^^
|
O^^
|
|
T. washingtonensis
|
CBS 544.50
|
?
|
B
|
?
|
B
|
B
|
?
|
B
|
|
Tilletiopsis spec. A
|
CBS111625
|
?
|
P
|
P
|
P
|
P
|
?
|
P
|
|
Tilletiopsis spec. A
|
CBS111628
|
7
|
?
|
?
|
?
|
?
|
?
|
?
|
|
Tilletiopsis spec. B
|
CBS111622
|
4’
|
O
|
O
|
O
|
O
|
O
|
O
|
|
Tilletiopsis spec. B
|
CBS111616
|
3
|
I
|
I
|
I
|
?
|
I^^
|
I^^
|
|
Tilletiopsis spec. B
|
CBS111617
|
3
|
I
|
I
|
I
|
I
|
I^^
|
I^^
|
|
Tilletiopsis spec. B
|
CBS111613
|
2
|
I
|
I*
|
I
|
I^^*
|
I^^
|
I^**^
|
|
Tilletiopsis spec. B
|
CBS111615
|
3
|
I
|
I
|
I
|
I^^*
|
I
|
I^^*
|
|
Tilletiopsis spec. B
|
CBS111608
|
3
|
I
|
I
|
I
|
I
|
I
|
I
|
|
Tilletiopsis spec. B
|
PD A10
|
3
|
I
|
I
|
I
|
I
|
I
|
I
|
|
Tilletiopsis spec. B
|
CBS111610
|
3
|
I
|
I
|
I
|
I*
|
I
|
I*
|
|
Tilletiopsis spec. B
|
CBS111611
|
3
|
I
|
I
|
I
|
I
|
I
|
I
|
|
Tilletiopsis spec. B
|
CBS111741
|
3
|
I
|
I
|
I
|
I
|
I
|
I
|
|
Tilletiopsis spec. B
|
CBS111620
|
3
|
I
|
I
|
I
|
I*
|
I^^
|
I*^^
|
|
Tilletiopsis spec. B
|
CBS111612
|
3
|
I
|
I
|
I
|
I
|
I
|
I
|
|
Tilletiopsis spec. B
|
CBS111609
|
3
|
I
|
I*
|
I
|
I*
|
I
|
I**
|
|
Tilletiopsis spec. B
|
CBS111619
|
3
|
I
|
I
|
I
|
I*
|
I^^
|
I*^^
|
|
Tilletiopsis spec. B
|
CBS111618
|
3
|
I
|
I
|
I
|
I*
|
I^^
|
I*^^
|
|
Tilletiopsis spec. B
|
CBS111614
|
3
|
I
|
I
|
I
|
I*
|
I
|
I*
|
|
Tilletiopsis spec. B
|
CBS111621
|
3
|
I
|
I
|
I
|
I
|
I^^
|
I^^
|
|
Tilletiopsis spec. C
|
CBS
111607
|
?
|
N
|
N
|
N
|
N
|
N
|
N
|
|
Tilletiopsis spec. D
|
CBS
111595
|
5
|
K
|
K
|
K
|
K
|
K
|
K
|
|
Tilletiopsis spec. D
|
CBS
111593
|
5
|
K
|
K
|
K
|
K
|
K
|
K
|
|
Tilletiopsis spec. D
|
CBS
111594
|
5
|
K
|
K
|
K
|
K
|
K
|
K
|
|
Tilletiopsis spec. E
|
CBS
111603
|
1
|
M
|
M
|
M
|
M
|
M
|
M
|
|
Tilletiopsis spec. E
|
CBS
111604
|
1
|
M
|
M
|
M
|
M
|
M
|
M
|
|
Tilletiopsis spec. F
|
CBS
111600
|
9
|
L
|
L
|
L
|
L
|
L
|
L
|
|
Tilletiopsis spec. F
|
CBS
111599
|
9
|
L
|
L
|
L
|
L
|
L
|
L
|
|
Tilletiopsis spec. F
|
CBS
111602
|
9
|
L
|
L
|
L
|
L
|
L
|
L
|
|
Tilletiopsis spec. F
|
CBS
111597
|
9
|
L
|
L
|
L
|
L
|
L
|
L
|
|
Tilletiopsis spec. F
|
CBS
111598
|
9
|
L
|
L
|
L
|
L
|
L
|
L
|
|
Tilletiopsis spec. F
|
CBS
111601
|
9
|
L
|
L
|
L
|
L
|
L
|
L
|
|
Tilletiopsis spec. F
|
CBS
111596
|
?
|
L
|
L
|
L
|
L
|
L
|
L
|
|
Tilletiopsis spec. F
|
CBS
111627
|
9
|
R
|
R
|
R
|
R
|
R
|
R
|
|
Tilletiopsis spec. G
|
CBS
111605
|
10
|
C
|
C
|
C
|
C*
|
C
|
C*
|
|
Tilletiopsis spec. G
|
CBS
111606
|
10
|
C
|
C*
|
C
|
C
|
C
|
C#
|