UKTC Archive

Re: Ganoderma applanatum/australe on n.maple - implications

Subject: Re: Ganoderma applanatum/australe on n.maple - implications
From: Annefrank Tree
Date: Dec 23 2011 17:15:00
Dear Gerrit,
 
You are correct my first language is German.
 
Your next project sounds very interesting to me. I have carried out field and 
glasshouse experiments utilising endo and ectomycorrhizal fungi, innoculated 
onto trees, in order to examine multitrophic interactions, biomass and inter 
specific fungal competition. I had not considered the time required to allow 
the tree "establish" before it could "afford" sugars. I will review my paper 
again, this may explain an unexpected result.
 
Regards
 
Freidrich
 

  

________________________________
 From: Viper Snake <snake24@xxxxx.nl>
To: UK Tree Care <uktc@xxxxxx.tree-care.info> 
Sent: Friday, 23 December 2011, 16:12
Subject: RE: Ganoderma applanatum/australe on n.maple - implications
 


Dear Freidrich,

I assume your native language is German, not Dutch like David assumes.

1) ... biotoof ...
1) In Dutch, it is always spelled biotroof.

2) I also find interesting the use of the term biotrophic parasite. I had not 
thought in these terms previously, my experience has been largely with 
mutualism and symbiosis. I have read however that such associations exist on 
a continuum of benefit depending largely alterations in environmental factors 
and may sometimes move towards parasitism. Though not proven it would be 
reasonably deduced that should the environmental conditions be restored 
mutualism would once more prevail. This may make an interesting area of 
research.
2) Next year, I'll start a research project (Wageningen University) on 
facultative ectomycorrhizal macrofungi such as Scleroderma citrinum, Paxillus 
involutus, Thelephora terrestris and Boletus badius, that do not seem to have 
lost their capacity to (temporarely) live as a saprotroph on and fruit from 
dead wood while "awaiting" to colonise the roots of a new partner and seem to 
refrain from retracting sugars for producing fruitbodies from the seedling 
and young tree partner until the tree can "stand on its own feet" by 
developing the foliage needed to produce enough energy (reserves) through 
photosynthesis to support itself and its tree species specific ecosystem, and 
can "afford" sharing the sugars with its symbionts, the mycelia need to fruit.

Regards,
Gerrit


Without wanting to complicate things any further, some citations on 
biotrophic and necrotrophic parasitic macrofungi from Scandinavian and Dutch 
literature and my own experience, I would like to have your opinion on, 
follow.

- The (biotrophic) parasitic Inonotus hispidus, that grows "on living 
hardwoods" and "dies with the tree" (my words), "is capable of killing 
sapwood (soft rot) in living trees" (Ryvarden & Gilbertson, European 
Polypores I).
- In my experience, the same thing applies to Inonotus cuticularis on Fagus 
and to Meripilus giganteus on Fagus and the 16 other tree species it is this 
far documented from, of which the mycelia of both species also invade and 
cause a soft rot of living tissues.
- The (biotrophic) parasitic Phellinus tremulae "spreads in the inner sapwood 
and inner wood of living trees in the absence of other organisms, indicating 
that it is a primairy parasite in aspen" (Wickström in : Ryvarden & 
Gilbertson, European Polypores II).
- And is Pholiota squarrosa obligate (biotrophic) parasitic or - according to 
Arnolds, et al. : Overzicht van de Paddestoelen in Nederland - necrotrophic 
parasitic, as it dies with the tree and never fruits from dead wood alone, 
which is very exceptional for parasitic Agaricales ?

So how would you label these macrofungi, as obligate (biotrophic) parasitic 
or necrotrophic parasitic ?


biotroof - food withdrawing to live parts of a host.

I think that this simply means that the parasite is obtaining its food from 
living tissues. This is okay as a literal translation of the original 
Greek: "bios" = life / "trophe" = food. I do not, however, think that this 
definition is precise enough to distinguish between a biotroph and a 
necrotroph (just as you implied in your previous message).

In Ainsworth and Bisby's Dictionary of the Fungi (8th edition, CAB 
International), the definition of biotroph is as follows:

"Biotroph: an obligate parasite growing on another organism, in intimate 
association with its cytoplasm."

I think that the Ainsworth & Bisby definition means almost the same as the 
one that I wrote yesterday. I did not, however, use the word "obligate" 
(meaning that the parasite can grow only on the living host; not on a 
non-living culture medium), because I am aware of a few rust fungi (for 
example) that have been grown on artificial media, with great skill and 
difficulty. On the other hand, it is quite useful to use the word 
'obligate', since this clearly shows that wood decay fungi cannot be 
biotrophic according to the strict definition. They must either be 
necrotrophic parasites (if they invade and kill living tissues) or 
saprotrophs (if they grow only in tissues that are already dead).

I use the definitions from this list (in Dutch) ( 
www.springerlink.com/content/tuj653216XXXXXXX ) and to add some more 
information on the subject of tree species specific strategies of parasitic 
macrofungi being part of and co-evolved within a tree species specific 
ecosystem (Keizer, 2007/2011), the following on the tree species colonised 
by Fomitopsis pinicola on the European continent.

I hadn't realised the use of "biotrophic" and "necrotrophic" were 
differently in the UK and on the continent; thanks for mentioning this.

I don't know whether other UKTC members share my unease about identifying 
species of Ganoderma in the field. I would feel happier if each species 
could be shown to have a distinct combination of characteristics (including 
hyphal structure, average spore-size, DNA-based criteria, thickness and 
hardness of the crust and the presence/absence of insect-galls). Perhaps we 
are wrong in the UK to be identifying a large proportion of the Ganoderma 
on beech as G. australe/adspersum. Various fungi do, however, show 
genuinely different frequencies of occurrence between the UK and parts of 
the near continent. For example, we seem to have more G. pfeifferi here. 
Also, Fomes fomentarius is much less frequent on beech (and on other tree 
species) in southern Britain than on the continent. It is more common in 
northern Britain, where the main host is birch.


-----Original Message-----
From: Viper Snake [mailto:snake24@xxxxx.nl]
Sent: 21 December 2011 19:33
To: UK Tree Care
Subject: RE: Ganoderma applanatum/australe on n.maple - implications




Dear David,

Although there is quite a difference in your definition of biotrophic 
and necrotrophic parasites and the definition of both terms on the 
continent, as is included in all Dutch, German and Scandinavian 
literature I refer to, I consider your definition to be the more 
precise.

And after always microscopically assessing what perennial species of 
Ganoderma I found on beech in The Netherlands and Germany, I have come 
to the conclusion, that about 95 % of perennial Ganoderma species on 
beech is G. lipsiense, followed by the rare G. pfeifferi and the 
extremely rare G. australe.

Regards,
Gerrit



From: d.lonsdale2@xxxxxxxxxxx.com
To: uktc@xxxxxx.tree-care.info
Subject: RE: Ganoderma applanatum/australe on n.maple - implications
Date: Wed, 21 Dec 2011 18:29:51 +0000

Dear Gerrit,

In answer to your question, I define the relevant terms as follows:

1. Biotrophic parasite: a parasite which obtains its nutrients from 
the living cells of the host (usually by the penetration of those 
cells, without killing them). Examples include rusts and mildews.

2. Necrotrophic parasite: a parasite which obtains its nutrients by 
killing cells of the host (usually by the secretion of enzymes and/or 
toxins). These parasites are usually able to grow also as saprotrophs 
(4, below) and they are therefore often alternatively called 
"facultative parasites". Examples include many fungi and bacteria 
that cause general dieback and/or decay of host tissue.

3. Hemibiotrophic parasite: a parasite which obtains its nutrients 
both as a biotroph (usually when it first penetrates host tissue) and 
as a nectroph (usually at a later stage, after an initial biotrophic 
phase). Examples include many organisms that cause leaf spot diseases.

4. Saprotroph: an organism which obtains its nutrients from the dead 
remains of one or more living organisms.

As far as I know, there are no wood decay fungi in categories (1) or 
(3) above. Some of them have the ability to grow into previously 
living sapwood, causing it to die (or become "dysfunctional" and then 
causing decay. I think that they can be regarded as necrotrophic 
parasites (2), but I do not like to use this term without 
qualification, since many of them live predominantly on wood that is 
already dead. This could be sapwood that has been damaged by injury, 
or it could be central wood or the tree, which has become heartwood 
or ripewood because of aging. 

Traditionally, wood decay fungi have been described as "parasitic" if 
they are found on living stems, branches or roots. I do not think 
that this is correct if the fungus concerned is colonising only wood 
that is already dead.

According to the above definitions, Ganoderma applanatum has been 
observed to be mostly saprotrophic, whereas G. adpsersum/australe has 
some capacity to act as a necrotrophic parasite. These observation 
seem to be confirmed by some experimental work by my friends Schwarze 
& Ferner at the University of Freiburg i. Br., Germany. (see: 
http://www.enspec.com/articles/ENSPEC%20Research%20Paper%20-%20Ganoderma%20on%20Trees.pdf
 )

Schwarze & Ferner found that G. adpsersum/australe was able to 
penetrate defensive barriers (reaction zones), thus growing into 
functional sapwood. It does not necessarily harm the tree seriously. 
Instead, it might be able to co-exist with the tree for many years, 
instead of dying out when it has utilised all the wood that was 
initially available to it. In some cases, however, the fungus does 
enough damage to the sapwood (especially in the roots of the tree) to 
cause the decline and perhaps death of the tree. Also, the decay can, 
in my experience, become overwhelmingly rapid if the wood becomes 
more aerated because of excessive pruning or the storm-breakage of 
major branches. 

As suggested in the recent correspondence, G. adpsersum/australe 
appears to behave differently in different host species. I think that 
it can be especially aggressive in species with which it has not 
co-evolved. I have seen examples where G. adspsersum/australe (or 
perhaps a similar-looking species of Ganoderma) seems to have killed 
exotic conifers such as Araucaria araucana. I agree that it can cause 
extensive decay in the broadleaved trees in your list. However, in 
Fagus sylvatica (one of its main hosts in the UK), the tree and the 
fungus often seem to co-exist for many years. The co-existence is 
probably even longer in species like Quercus robur and Q. petraea, 
which have durable heartwood and therefore tend to become decayed 
more slowly.

Regards,
David

-----Original Message-----
From: Viper Snake [mailto:snake24@xxxxx.nl]
Sent: 21 December 2011 14:14
To: UK Tree Care
Subject: RE: Ganoderma applanatum/australe on n.maple - implications




David,

1). Tony Croft has been using the term 'biotrophic parasite' in 
relation to G. adspersum/australe on here. I questioned him about 
these labels and asked him whether he could point me to any research 
that demonstrated that the fungus was 'biotrophic' and/or 
'parasitic', because this was news to me, but he hasn't replied yet. 
Can you point me in the right direction?

1). Tony uses my terms and the results of my in situ research on 
biotrophic and/or necrotrophic parasites, which I already have 
explained on Arbtalk, see : 2. 
http://arbtalk.co.uk/forum/529703-post53.html .

2). Similarly, that G. adspersum/australe is deterministically fatal 
to Acers, no matter the circumstances, is also news to me. Where does 
the research for this conclusion come from ? 


2). From my own field research on the effects on the stability and 
condition of different deciduous tree species of the biotrophic 
parasitic G. australe, of which the mycelium causes a white rot with 
selective delignification, that is most detrimental to Acer, 
Platanus, Populus, Salix, Tilia, Aesculus (Anne Frank tree) and 
Quercus rubra.

Gerrit





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