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Introduction
I am an evolutionary ecologist, writer and artist. I was born in Nairobi, Kenya in 1950 and gained BA and PhD degrees at King’s College, Cambridge in 1972 and 1975. I was a Reader in Biology at the University of Bath from 1985 to 2011 and have published numerous papers and books, the latter including, most recently, 'The Origin of Life Patterns in the Natural Inclusion of Space in Flux’. I was President of the British Mycological Society in 1998 and of Bath Natural History Society from 2012 - 2018. Since 2000, I have been pioneering understanding of 'natural inclusion', the receptive-responsive evolutionary relationship between space and energy in all material form. See http://www.spanglefish.com/exploringnaturalinclusion/
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Publications (148)
This chapter draws on my experience of presenting a trans-disciplinary course, entitled ‘Life, Environment and People’, at the University of Bath, UK, from 2001 to 2011. The course was available to final year undergraduate students studying for degrees in biological sciences, natural sciences, management and psychology. It was based on my understan...
The age-old contradiction between competition and cooperation as alternative perceptions of life and evolution is reconciled by the γ-principle of natural inclusion. This principle has three fundamental tenets, all of which need to be recognised if it is to be understood comprehensively: (1) Natural bodily boundaries are intrinsically dynamic, not...
For millennia, the Western mindset has been predisposed to the inherited custom to split mind and matter, emotion and cognition, art and science, the spiritual and the intellectual, the inner and outer, contemplation and objective inquiry. Our contemporary sense of dislocation from nature has arisen from this objectivistic perception. The current s...
Drawing on personal experience of early childhood in Kenya and later life in the UK, a contrast is recognised between culturally embedded attitudes of mind that either categorically divide or totally unite human identity from or with its natural neighbourhood. Neither of these attitudes makes consistent natural sense, and both can result in false a...
This pioneering article explores why, fundamentally, so many of us abide by a perception that objectively isolates our receptive self-centres as singular points of mass, and asks whether there is a way of perceiving Nature and human nature that incorporates rather excludes our emotional, soulful and spiritual qualities. The answers offered come dow...
Abstract geometries, whether Euclidean, non-Euclidean or fractal, cannot realistically describe the dynamic nature of life patterns. This is because they all depend on the unnatural dissociation of matter from space, as if matter and space are independent from each other. This leads to either a reduction of particles into dimensionless point masses...
The receptive presence of intangible space is central to self-identity and love and understanding of one another. It is in the heart of our ‘Being’ as sentient life forms, where it combines with energetic flux to co-create tangible natural bodies as continuous ‘Becomings’. If it were to be personified, she could be known as Grace, that dwelling pla...
The author’s early life experiences that contributed to his appreciation of his own and others’ self-identities as receptive centres of awareness are briefly described. This appreciation is contradicted by objective perceptions of reality that exclude individual receptivity from consideration by mentally isolating natural bodies within definitive b...
There is abundant direct and indirect evidence that life and the cosmos have not remained unchanged since an act of instantaneous creation. Life and the cosmos evolve through an ongoing cumulative process of energetic transformation that is evident even within the life spans of individual organisms. Efforts to understand this process have, however,...
Careful study reveals a variety of recurrent natural patterns of life that are evident over vastly different scales of organization, within living cells, multicellular organisms, colonies, populations, communities and ecosystems: stars and stripes, circles, lines, spirals, rivers, ripples and crazy paving. The vital role of receptive perception and...
Extraordinarily varied patterns of structure, function and relationship emerge and co-evolve over nested scales of organization in living systems. These patterns recur in cells, individuals, species and ecosystems. Their evolution depends on the generation and sustainability of diversity, not its progressive elimination in favour of what is special...
Abstract and natural perceptions of life patterns affect human relationships and social organization in different ways. Abstract thinking is prone to isolate humanity from its natural source of life and love, through its disregard of individual receptivity. This is the root cause of the estrangement from one another and our natural neighbourhood th...
Understanding the relationship between human cultural psychology and the evolutionary ecology of living systems is currently limited by abstract perceptions of space and boundaries as sources of definitive discontinuity. This Brief explores the new understandings possible when space and boundaries are perceived instead as sources of receptive conti...
A new understanding of biology shows that if pre-cellular life originated as a community effort, then individuality could have evolved as a result of working together.
Trying to identify the smallest genome of independently living cells reveals a curious problem: Treating organisms as if they are self-contained entities, isolated from their neighborhood, is a profound mistake. All organisms depend on their environment for energy, carbon, and mineral nutrients to grow and reproduce. No plant, animal, or microbe ca...
This contribution to the Common Knowledge symposium “Fuzzy Studies” argues that the inclusion of space in form brings varying degrees of fuzziness and fluidity to all natural identities. Such inclusion is vital to evolutionary creativity, from subatomic to cosmic scales of natural energy flow. Examples abound throughout the natural world of indeter...
Fungal mycelia can alter their organizational pattern in such ways as to produce alternative phenotypes. The latter allow mycelia to explore for, assimilate, conserve, and redistribute resources in spatially and temporally heterogeneous niches. It is suggested that mycelia produce alternative phenotypes by operating as nonlinear (feedback regulated...
Psychology is not alone in its struggle with conceptualizing the dynamic relationship between space and individual or collective identity. This general epistemological issue haunts biology where it has a specific focus in evolutionary arguments. It arises because of the incompatibility between definitive logical systems of 'contradiction or unity',...
Learning is an evolutionary process that enables us to develop the skills necessary to sustain our lives in a complex, changing world. The way that educators understand this process is liable to affect their practice in very fundamental ways. Where they follow an 'adaptive' evolutionary model, their practice is likely to be highly prescriptive, wit...
Isolates of the mycelial cord-forming agaric, Tricholomopsis platyphylla, were obtained from an oak stand and their mating and mycelial interactions studied. The fungus was found to have tetrapolar incompatibility and genetically different dikaryons were strongly mutually antagonistic in culture. On the basis of such antagonism, 22 mycelial types,...
Experimental systems for studying the migration of nuclei into mating-type compatible homokaryons across a pre-existing or developing somatic incompatibility barrier are described. Using these systems with Stereum hirsutum Willd. ex Fr. it was found that different donor nuclei migrated from implants at more or less equal rates through acceptor homo...
Substantial mycelial cord systems of Hypholoma fasciculare (Huds. ex Fr.) Kummer, Phallus impudicus (L.) Pers., Phanerochaete (Ph.) velutina (DC ex Pers.) Parmasto, Phanerochaete laevis (Fr.) Erikss. & Ryv. and Steccherinum fimbriatum (Pers. ex Fr.) Erikss. had developed 2 years alter direct inoculation into the soil and litter of a range of woodla...
Almost 1000 wood blocks, 8 cm3, of beech (Fagus sylvatica L.), colonized by the mycelial cord-forming basidiomycetes Hypholoma fasciculare (Huds. ex Fr.) Kummer., Phallus impudius (L.) Pers., Phanerochaete (Ph.) laevis (Fr.) Erikss & Ryv., Ph. Velutina (DC ex Pers.) Parmasto, Steccherimum fimbriatum (Pers. ex Fr.) Erikss. and Tricholomopsis platyph...
Focuses on the problems facing an individual, plant-infecting fungal mycelial agent in terms of the maintenance of evolutionary fitness, both of itself and of its progeny. The extent to which competition with other genets of the same or different species is a major determinant of its life style, and depends largely on the degree to which it is adap...
The ciid beetles Octotemnus glabriculus and Cis boleti exploit different developmental stages of fruit bodies of their preferred host fungus Coriolus versicolor. Larvae of the smaller beetle, O. glabriculus, mainly use young, expanding, fruit bodies; adults of O. glabriculus are predominantly found in young fruit bodies. By contrast, adults and lar...
For centuries, our understanding of how we relate to our environment has been impeded by the deliberate exclusion of context which comes from rationalistic modes of enquiry that place unrealistically discrete boundaries between ‘insides’ and ‘outsides’, ‘subjects’ and ‘objects’ and ‘self’ and ‘other’. Now that the global impact of human technology...
The mechanism of thatch deterioration is poorly understood Preliminary observations were made on 16 roofs, all thatched with combed wheat reed. Thatch wears away from the exposed surface of the roof. Three distinct zones (the outer, middle and inner) develop and appear to represent three stages in the decay of thatch. The inner zone is a region of...
On the basis of the evidence that insect fungivory has the potential to affect fungal reproductive fitness, we investigated the effects of two specialist ciid beetles (Octotemnus glabriculus and Cis boleti) on the reproductive potential of their host fungus, Coriolus versicolor. We found, from field data, a negative correlation between the number o...
Most ciids (Ciidae) are strict fungivores specialized on fruit bodies of wood-rotting fungi. The Ciidae includes both specialist and generalist species. Recent evidence suggests that ciids locate and discriminate their potential hosts based mainly on fungal odours. In this study, we investigated the field distribution of ciids in a local woodland n...
The mechanism of thatch deterioration is poorly understood. Preliminary observations were made on 16 roofs, all thatched with combed wheat reed. Thatch wears away from the exposed surface of the roof. Three distinct zones (the outer, middle and inner) develop and appear to represent three stages in the decay of thatch. The inner zone is a region of...
Fungi are amongst the simplest of eukaryotes. Their study has provided useful paradigms for processes that are fundamental to the way in which higher cells grow, divide, establish form and shape, and communicate with one another. The majority of work has been carried out on the budding yeast Saccharomyces cerevisiae, but in nature unicellular fungi...
The way that organisms respond to one another and their environment in natural populations depends critically on the dynamic processes that produce and limit phenotypic diversity. However, the fundamental nature of these processes, and the extent to which they are genetically prescribed, is arguably the most challenging issue facing evolutionary bi...
Perhaps like many other mycologists, my initial encounters
with fungi filled me with a desire to engage with mysteries:
mysteries embedded in the diversity of form and function in
fungi and how this diversity might be related to the life of the
forest in which so many of these organisms appear to find
their homes. In all good mysteries, the re...
Under a variety of conditions, the hyphal density within the expanding outer edge of growing fungal mycelia can be spatially
heterogeneous or nearly uniform. We conduct an analysis of a system of reaction-diffusion equations used to model the growth
of fungal mycelia and the subsequent development of macroscopic patterns produced by differing hyph...
Fungal mycelia often epitomise the growth and pattern-generating properties of a wide variety of indeterminate living systems. They may, therefore, provide an experimentally accessible model for the study of other networked structures such as nervous and vascular systems. In the following, we use a system of coupled reaction-diffusion equations to...
Fungal mycelia epitomize, at the cellular level of organization, the
growth and pattern-generating properties of a wide variety of
indeterminate (indefinitely expandable) living systems. Some of the more
important of these properties arise from the capacity of an initially
dendritic system of protoplasm filled, apically extending hyphal tubes
to an...
Twelve heterokaryotic strains of Heterobasidion annosum (Fr.) Bref. containing nuclei and mitochondria derived from British and North European populations were prepared by pairing homokaryotic strains and isolating hyphal tips from either side of interaction interfaces. Conidia derived from the heterokaryons had high germinability and were predomin...
Chaos theory deals with systems whose long-term behaviour or output tends to be complex, irregular, sensitive to small changes in initial conditions and unpredictable at particular localities. Non-linear theory is argued to provide a basis for understanding the complexity, interconnectedness and limits to the predictability of natural patterns of d...
As energy-using systems, all life forms must possess boundaries. However, these boundaries can neither be fully open nor fully
closed to energy transfer if they are not to become infinitely dispersed or forever static. Instead, they define the reactive
interfaces between the insides and outsides — i.e. the “dynamic contexts” — of living systems. Th...
summaryRatios of nuclear genotypes observed in conidia from heterokaryotic strains of Heterobasidion annosum (Fr.) Bref., obtained from pairings between sympatrically derived, sib-related and non-sib-related homokaryons, commonly deviated from 1:1. Ratios were temporally stable, and the genotypes examined could be ranked in a strict dominance hiera...
Fungal mycelia are capable of growth and persistence over extended, potentially unlimited periods (see also Carlile, Chapter
1). In this respect they contrast with determinate body forms, such as those of many animals and unicellular organisms, which
have pre-set limits in space and time. Moreover, as different parts of a mycelial boundary expand,...
An understanding of sources of phenotypic variation in natural ascomycete populations is critical to the evaluation of criteria for taxonomic delimitation. Such variation may reflect genetic differences between individuals due to mutational and recombinatorial processes, or result from epigenetic changes in developmental patterns within heterogeneo...
The interrelationships between trees, and the fungi that inhabit and decay their xylem, are dynamic, complex and difficult to analyse into simple sequences of cause and effect. This is because both participants function, and interact, as highly versatile, open-ended or “indeterminate” hydrodynamic (“self-plumbing”) systems. As such they are capable...
Strains of the basidiomycete fungi Stereum hirsutum from the USSR and Stereum complicatum from the eastern USA were interfertile, yielding viable, variable, meiotic progeny. However, interactions between paired homokaryotic mycelia of these species were unstable, yielding a uniform mycelial mat on the S. complicatum side, but a macroscopically hete...
Three separate collections were made in southern England of Phanerochaete magnoliae forming hydnoid hymenial surfaces through the tubes of Datronia mollis basidiomes. Isolates of P. magnoliae exhibited unusual patterns of development and interaction with other fungi. Germinating basidiospores produced an extensive, sparsely branched, rapidly extend...
Hyphal and mycelial interactions within progeny sets of single basidiospore isolates derived from Heterobasidion annosum fruit bodies from various parts of the northern hemisphere were studied microscopically using microculture chambers and with pairing tests, respectively. The behaviour within different sets was highly variable with respect to mat...
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The mycelium of higher fungi is portrayed as a developmentally versatile collective in which an initially dendritic pattern of branching is converted, by hyphal anastomosis, into a communication network. Spatial and functional patterns in the mycelium depend on the establishment of gradients allowing flow of protoplasmic resources and organelles vi...
The mycelium of higher fungi is portrayed as a developmentally versatile collective in which an initially dendritic pattern of branching is converted, by hyphal anastomosis, into a communication network. Spatial and functional patterns in the mycelium depend on the establishment of gradients allowing flow of protoplasmic resources and organelles vi...
Pairings between certain combinations of homokaryotic strains of Stereum spp. give rise, following immigration of non-self nuclei, to extensive zones of appressed, degenerative mycelium. Aerial, crystalline filaments, up to 35 m wide and ⩾ 6 cm long, emerge from this mycelium at localized foci which appear to act as sinks for mycelial resources. Th...
Outcrossing populations of the Coniophora puteana complex had a multiallelic unifactorial (bipolar) homogenic incompatibility (mating) system regulating the emergence of a secondary mycelium (stable mating-type heterokaryon) between paired single basidiospore-derived strains (homokaryons). Emergence of the secondary mycelium involved a switch to fa...
Populations of the wood-decaying basidiomycete, Stereum hirsutum (Willd. ex Fr.) S. F. Gray, from different geographical regions, reproduce either by outcrossing or non-outcrossing. Outcrossing generates variable progeny whilst non-outcrossing results in clonal sub-populations.
Pairings between single-basidiospore-derived strains from non-outcrossi...
Individual mycelial genotypes (genets) of Hymenochaete corrugata are able to traverse the canopies of coppice-grown hazel, Corylus avellana, by producing sclerotized mycelial pads which bridge between and bind together contiguous stems from the same and different stools. The pads may also allow attachment to other trees, e.g. elm (Ulmus sp.), climb...
Pairings between strains of Stereum hirsutum (Willd. ex Fr.) S. F. Gray pp 2% malt agar were analysed using changes in mycelial morphology and non-self recognition properties as evidence of genetic transfer between the participants. In some combinations the morphological evidence suggested the- uniform emergence of a heterokaryon following mating a...
Cultural studies of the forest root pathogen Heterobasidion annosum (Fr.) Bref. have shown that the mycelium of the fungus possesses a high degree of developmental versatility, being able to grow in a variety of distinctive functional modes. Consequently the fungus can alternate between conidiogenous and meiotic reproductive pathways, rapid, sparse...
Evidence of a non-outcrossing breeding strategy, resulting in production of morphologically indistinguishable, somatically compatible ascospore progeny from individual perithecia, was detected in natural populations of Hypoxylon multiforme and an undescribed taxon provisionally designated ‘H. purpureum’. However, in both cases the populations were...
Evidence of outcrossing was detected in populations of Hypoxylon fragiforme, H. fuscum, H. mammatum, H. nummularium H. rubiginosum and H. serpens. Single ascospore-derived strains from the same perithecium mostly produced demarcation zones when paired against one another on 2% malt agar, as did all wood or single ascospore strains obtained from dif...
Cord systems of Hypholoma fasciculare, Phallus impudicus, Phanerochaete laevis, Phanerochaete velutina and Steccherinum fimbriatum developed progressively in non-sterile soil from advancing mycelial fronts within which linear aggregation of hyphae was followed by lysis of diffuse mycelium.Extension rates of the cord systems were generally linear an...
Living hyphae of Stereum hirstutum behavaved similarly prior to fusion on a cellophane membrane whether they were genetically the same or different. Fusions were generally initiated within 5-10 h of mycelial interdigitation. A period (10-35 min) of interfacial expansion usually preceded opening of the fusion pore, but in certain combinations involv...
The extension rates of Clitocybe nebularis (Batsch ex Fr.) Kummer strains on 2% malt agar were only 30–40%, of those, up to 3.4 mm d−1, observed in woodland at equivalent exponential mean temperatures. Extension of mature field systems was accomplished by mycelial annuli or arcs 30-40 cm wide, differentiated into a leading edge of mycelial cords fo...
Inoculum blocks of wood colonized by the basidiomycetes Hypholoma fasciculare (Huds.: Fr.) Kummer and Phanerochaete velutina (DC: Pers.) Parmasto were placed in plastic trays containing moist (matric potential −0.007 MPa), unsterilized soil. When ‘baits’, in the form of beech (Fagus sylvatica L.) wood blocks, descaled pine (Primus sylvestris L.) co...
Inoculum blocks of wood colonized by the basidiomycetes Hypholoma fasciculare and Phanerochaete velutina were placed in plastic trays containing moist, unsterilized soil. When beech Fagus sylvatica wood blocks, descaled pine Pinus sylvestris cones, beech twigs, beech leaves or pine needles were introduced, H. fasciculare consistently responded by r...
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Wood blocks colonised by the basidiomycetes Phallus impudicus, Phanerochaete laevis and Steccherinum fimbriatum were placed individually in plastic trays containing moist, unsterilised soil. All three fungi grew out radially from the inoculum blocks, forming networks of mycelial cords. Outgrowth patterns of P. impudicus and P. laevis were similar i...
The mycelial interactions of strains of six cord-forming wood-inhabiting basidiomycetes were studied both against each other and against other fungi including Armillaria species on 2% malt extract agar, in wood lengths, and in non-sterile soil. Generally, cord-formers could be ranked in a combative order Phanerochaete velutina (DC ex Pers.) Parmast...
Rosellinia desmazieresii was isolated from 11 disease rings in 5 different sand dune slacks at Ainsdale National Nature Reserve, Lancashire. Single ascospore isolates from the same perithecium, or wood- or ascospore-derived isolates from the same ring, were somatically compatible and morphologically identical, indicating a non-outcrossing breeding...
Synopsis
Some roofs that are thatched with combed wheat reed develop patches of more rapid decay within an otherwise uniform matrix of less rapid decay. Changes in the length and weight of reed from patches and matrix were estimated for one roof by comparison with samples of unused, undecayed reed. The fungal communities associated with patches and...
Nowhere, we believe, can the presence of fungal communities, their structure, dynamics, and diversity, be more explicit and susceptible to direct analysis than in decaying wood. In consequence, wood provides an excellent venue, both for the study of community interactions, and for the development of a conceptual framework within which they can be r...
Despite their different fundamental organization, ant colonies and mycelia of fungi exhibit striking similarities in their social organization. Both are collectives of genetically related or identical semi-autonomous units, consisting respectively of discrete multicellular individuals and hyphae. There is a variety of parallels in their foraging an...
Temporary parasitism by individual of the wood-decaying basidiomycete Lenzites betulina allows them to gain selective access to wood occupied by populations of pioneer basidiomycete colonisers in the genus Coriolus. Pseudotrametes gibbosa is similarly temporarily parasitic on species of Bjerkandera. This strategy facilitates possession by the paras...
Genetically different isolates of Daldinia concentrica always produced demarcation zones when paired against one another. The morphology of these zones varied with respect to the width and shape of regions of white aerial mycelium (wam).
In pairings between single ascospore isolates from the same perithecium, evidence was obtained for a multigenic...
Mycelia of Hymenochaete corrugata exhibited two distinctive developmental patterns when cultivated on 2 % malt agar, resulting in either an appressed, yellow-brown pigmented colony or a white colony with extensive aerial mycelium. The switch from one colony type to the other sometimes occurred spontaneously, resulting in formation of sectors. On ot...
Before fusion, hyphae of the basidiomycete Phanerochaete velutina responded similarly to one another when grown on a cellophane membrane, regardless of whether they were genetically the same or different. Long-range (up to 250 μm) curvature (homing) to specific sites in the lateral wall of recipient compartments often occurred in fusions involving...
~~~ ~~~ Wood blocks colonized by the basidiomycetes Hypholoma fasciculare and Phanerochaete velutina were placed in plastic trays containing moist unsterilized soil. Both fungi grew out radially from the inoculum blocks in the form of networks of mycelial cords. When a second, uncolonized wood block, or set of wood blocks, was provided as a ‘bait’...
Various fully compatible and hemi-compatible pairings were made on 2 % malt agar between laboratory-synthesized heterokaryons and homokaryons of British collections of Stereum hirsutum. In all cases heterokaryons arose within the originally homokaryotic mycelia. However, the patterns of interaction between the colonies, and the number and genotypes...
More than 450 beech (Fagus sylvatica L.) logs, 10 to 20 cm in diameter and 30 to 40 cm long, were cut from freshly felled trees and placed upright 1 m apart with their bases buried up to 10 cm deep in the ground in a plot of about 600 m2 in a mixed deciduous woodland in the Forest of Dean, Gloucestershire, UK. Fungal colonization via the aerial and...
The spatial development, in beech logs exposed to air-borne and soil-borne inoculum, of mature fungal communities containing mutually or unilaterally exclusive mycelia of decay species is described and related to the ecological strategies and interaction of participant individuals. A combative heirarchy is recognized between (i) ruderal and/or stre...
Patterns of fungal colonization from the base of cut beech logs placed upright and partly buried in the ground at a mixed deciduous woodland site were fundamentally different from those from the aerial cut surface.
Principal colonists resulting in decay, and mostly arriving within three months, were the ascomycete Xylaria hypoxylon (L. ex Hooker) G...
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