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Chestnuts: History
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Subjects: Plant Sciences
Contributor: GABRIELE BECCARO

Chestnuts (Castanea spp.) are trees of relevant economic interest in the agro-sylvicultural contexts of mountain regions throughout the temperate zone, particularly in the northern hemisphere.

  • Castanea spp.
  • defensive mutualism
  • endophytes

1. Introduction

Chestnuts (Castanea spp.) are geographically distributed in three main areas throughout the world (Asia, Europe, North America) where they have an invaluable cultural heritage but also an important economic and environmental role in many agroforestry systems. Their nuts provided for centuries a dietary staple in rural areas and, when dried, a stored food for the whole year; the wood is still today used as firewood or building timber.
The genus Castanea (x = 12, 2n = 24), chestnuts and chinkapins, belongs to the Fagaceae (Cupuliferae) family, which includes 6 genera (Castanea, Quercus, Castanopsis, Fagus, Nothofagus, Lithocarpus) and approximately 1000 species. It is closely related to the genus Castanopsis [1], is widespread in the boreal hemisphere and encompasses 13 species. The natural distribution of the European or sweet chestnut (C. sativa) covers Europe and some Mediterranean countries. In China, Japan, Vietnam, North and South Korea C. mollissima, C. crenata, C. seguinii, C. henryi occur. In North America, C. dentata is still present only along the Appalachian Mountains, due to the devastating impact of the chestnut blight. Castanea pumila is found in the southeastern United States [2][3].
Castanea species show different ecological and morphological traits, vegetative habit, wood and nut characteristics, pedoclimatic adaptability, resilience and resistance to biotic and abiotic stresses, reflecting the adaptation of this genus to different environmental conditions [4]. The main cultivated species are C. sativa, C. mollissima and C. crenata, due to their large nut size, and marroni cultivars (C. sativa) are considered the most commercially valuable. Castanea sativa and C. dentata are also cultivated for timber production, whilst many interspecific hybrids are used for nut production or as rootstocks [5].

2. Ecological and Technical Features of Chestnut Orchards

The ecological features of chestnut orchards, their agronomic management and architecture strongly depend on species and country. The best soils for chestnut are deep, soft, volcanic in origin, and rich in phosphorus, potassium, and organic matter. The soil pH should be in the range 5.0−6.5; therefore, soils with active limestone are not optimal, because Castanea species are very sensitive to high pH values. Soil permeability is very important. In fact, the crop performs better in well-drained, loam to sandy-loam soils, while heavy, washed out, clayey, stagnant soils which favor root rot must be avoided [6].
Chestnut tolerates cold winters and requires average temperature of 8–15 °C, with a monthly average of 10 °C for at least six months. Castanea sativa is more cold-resistant (−15 to −20 °C) than the Euro-Japanese hybrids. Despite late bud-break (March–April in the Northern Hemisphere), the trees may be prone to spring frosts which damage young shoots. Temperatures of 27−30 °C are necessary during pollination. European and Japanese cultivars require about 800–900 mm/year of rainfall, well distributed during the growing season, while Euro-Japanese hybrids and C. mollissima are much more waterdemanding (1200–1300 mm/year). In temperate climates, sweet chestnut should not be planted above 800–900 m, while for hybrids the maximum altitude is about 500–600 m [5].
Ecological features of a chestnut orchard can be influenced by many natural and anthropogenic parameters, such as soil, climate, rootstock, cultivar, cover crops, irrigation system and fertilization [7]. Concerning soil, an increase in the organic matter content is often obtained through sheep and cattle pasture, outside the harvesting period, or with the use of manure (10–15 tons/ha/year). Mown grass, leaves, husks and the small pruned material are often left on the ground as they provide further enrichment in organic matter. Fertilization of the orchard is every year carried out according to soil conditions and uptakes [8]. Biological or conventional management are both adopted by chestnut growers and could strongly influence the crop ecological features.
The orchard model, traditional or intensive, could also strongly influence the crop ecological features. Traditional orchards are mainly located in mountainous areas while intensive orchards are a relatively recent practice and are usually located in lowlands, quite far from the typical ecological conditions of the species [9]. For all species, except in traditional European chestnut orchards, the general trend is to increase plant density to develop in a relatively short time maximum bearing per unit area. Plantation density can affect the orchard microclimatic conditions and can range from 100 to 550 trees/ha, based on species, variety, genotype-environment interactions and cultural practices. For traditional plantations of C. sativa, spacing ranges from 8–10 m apart in rows and 8–12 m between rows. For the most vigorous Euro-Japanese hybrids the distances range between 7 × 8 m (178 trees/ha) and 8 × 10 m (125 trees/ha). Castanea sativa and the Euro-Japanese hybrids can be cultivated in high density plantations (3 m × 10 m). For C. crenata distances of about 5 × 7 m (285 trees/ha, in deep fertile soils) or 7 × 7 m (204 trees/ha) are recommended. Castanea mollissima, grown in China and the Unites States, is managed in a high or semi-high density scheme, due to the smaller tree size. Planting patterns may be rectangular, squared or triangular, but the first scheme is mostly used because of easier management [3]. Nowadays, many traditional plantations need renewal and recovery after years of abandonment or following the attacks of pests and diseases that have compromised their efficiency.

3. Occurrence of Endophytic Fungi in Chestnuts and Ecological Implications

Although characterized by synchronized development inside the plant tissues and nutrient transfer at the interfaces [10], EMF are not included in the category of endophytes since at some extent they also grow saprophytically in the soil. Conversely, the definition of endophytes, on which this paper is more specifically centered, does not reflect a specialized nutritional function, and it is conventionally applied to microorganisms that colonize living, internal tissues of plants without causing any immediate, overt negative effect [11].
Data concerning occurrence of endophytic fungi of chestnuts (Table 1) are only available from a low number of countries, indicating that consideration for the ecological implications and the economic impact related to this component of biodiversity is still quite limited. Possibly connected with a higher importance as an economic crop in Europe, most investigations have been carried out on C. sativa, with 76 taxa reported so far (2/3 of which identified at the species level), while the species C. crenata, C. dentata and C. mollissima appear to have been less frequently investigated, basically in Japan, the United States and China. As a general aspect, identifications concerning stem (shoots, branches, etc.) refer to the occurrence of endophytes in subcortical tissues, while no findings were reported from xylem [12][13].
In some cases, conditions of detection did not fully meet the basic requirements referring to the true endophytic condition. In fact, a recent study based on ITS meta-barcoding was addressed to establish whether fungal communities within cynipid galls are different from foliar endophytes. Unfortunately, results were basically presented with reference to classes to which the detected OTUs belong, and identifications at the species level was only carried out for the main OTUs, without distinguishing their origin (gall or leaf) [14]. Therefore, entries in Table 1 referring to this study must be taken with circumspection. Moreover, some uncertainty is entailed in a few reports concerning endophytic fungi associated with cankers produced by C. parasitica on both C. sativa and C. dentata [15][16][17][18][19][20]. Even if in these cases the plant material used for isolations was not asymptomatic following infection by a known pathogen, the sterilization procedure ensured that at least the isolated fungi had colonized the plant tissues before sampling and were not epiphytic contaminants.
Table 1. Endophytic fungi reported from Castanea spp.
Endophyte 1 Plant Part Country Reference
Castanea sativa
Acremonium cf. curvulum shoot (phellem) Bellinzona, Switzerland [12]
Alternaria alternata several plant parts Eurobin and Monbulk, Australia [21]
shoot Ticino, Switzerland [22]
bud Ankara, Turkey [23]
bark Vinhais, Portugal [24]
Alternaria sp. shoot Geneve and Ticino, Switzerland [25]
leaf or gall Southern Tuscany, Italy [14]
bud Ankara, Turkey [23]
Apiognomonia errabunda leaf Cureglia and Zarei, Switzerland [26]
Arcopilus aureus * bark Vinhais, Portugal [24]
Arthrinium arundinis leaf Vejoris, Spain [27]
Aspergillus sp. stem Black Sea region, Turkey [15]
bud Ankara, Turkey [23]
Asterosporium sp. shoot (phellem) Bellinzona, Switzerland [12]
Aureobasidium pullulans shoot Ticino, Switzerland [22][25]
Aureobasidium sp. bud Ankara, Turkey [23]
Biscogniauxia mediterranea bark Valpaços and Vinhais, Portugal [24]
Botryosphaeria dothidea shoot Ticino, Switzerland [25]
leaf or gall Southern Tuscany, Italy [14]
Botryosphaeria sp. several plant parts Eurobin and Monbulk, Australia [21]
Botryotinia pelargonii leaf or gall Southern Tuscany, Italy [14]
Botrytis cinerea several plant parts Eurobin and Monbulk, Australia [21]
Chaetomium sp. several plant parts Eurobin and Monbulk, Australia [21]
Cladosporium cladosporioides several plant parts Eurobin and Monbulk, Australia [21]
Cladosporium sp. fruit Eurobin and Monbulk, Australia [21]
bud Ankara, Turkey [23]
Colletotrichum acutatum shoot (phellem) Bellinzona, Switzerland [12]
leaf, shoot Monti Cimini, Italy [28]
leaf Vejoris, Spain [27]
Coprinellus domesticus bark Oghuz, Azerbaijan [17]
Coryneum modonium shoot (phellem) Bellinzona and Murg, Switzerland [12]
Cryphonectria parasitica shoot (phellem) Bellinzona, Switzerland [12]
sprout Fossemagne, France [29]
bark Valpaços and Vinhais, Portugal [24]
Cytospora chrysosperma bark Valpaços, Portugal [24]
Cytospora diatyrpelloidea bark Valpaços and Vinhais, Portugal [24]
Cytospora eucalypticola bark Valpaços, Portugal [24]
Cytospora quercicola bark Vinhais, Portugal [24]
Dendrostoma castaneum * shoot (phellem) Bellinzona and Murg, Switzerland [12]
fruit Toricella, Switzerland [30]
branch Astroni Nature Reserve, Italy [31]
Diaporthe amygdali bark Oghuz, Azerbaijan [17]
Diaporthe eres shoot Geneve and Ticino, Switzerland [22][25]
Diaporthe foeniculina branch Astroni Nature Reserve, Italy [32]
Diaporthe sp. shoot (phellem) Bellinzona and Murg, Switzerland [12]
leaf or gall Southern Tuscany, Italy [14]
bud Ankara, Turkey [23]
Diplodia seriata leaf or gall Southern Tuscany, Italy [14]
Diplodina castaneae * shoot (phellem) Bellinzona and Murg, Switzerland [12]
stem, twig Chablais and Ticino, Switzerland [32]
stem Northern Spain [32]
bark, stem Ismailly, Qabala, Sheki (Azerbaijan) [17]
Epicoccum nigrum several plant parts Eurobin and Monbulk, Australia [21]
bark Balakan, Azerbaijan [17]
bud Ankara, Turkey [23]
Eutypella sp. bark Zagatala, Azerbaijan [17]
Fusarium ciliatum leaf or gall Southern Tuscany, Italy [14]
Fusarium lateritium leaf or gall Southern Tuscany, Italy [14]
Fusarium oxysporum leaf or gall Southern Tuscany, Italy [14]
Fusarium sp. fruit Eurobin and Monbulk, Australia [21]
leaf or gall Southern Tuscany, Italy [14]
leaf Vejoris, Spain [27]
bud Ankara, Turkey [23]
Gnomoniopsis castaneae * several plant parts Eurobin and Monbulk, Australia [21]
bark, flower, leaf several locations in New Zealand [13]
several plant parts Cuneo province, Italy [33][34]
flower, leaf, stem Southern Australia [35]
shoot several locations in Northern Italy [36]
fruit several locations in Switzerland [37]
buds Aosta Valley and Piedmont, Italy [38]
shoot Geneve and Ticino, Switzerland [22][25]
several plant parts Monti Cimini, Italy [28][39]
leaf or gall Southern Tuscany, Italy [14]
leaf Vejoris, Spain [27]
leaf Netherlands [40]
branch Astroni Nature Reserve, Italy [31]
Hyphodermella rosae bark Ismailly and Shaki, Azerbaijan [17]
Hypoxylon fragiforme shoot (phellem) Bellinzona and Murg, Switzerland [12]
Irpex lacteus bark Balakan, Azerbaijan [17]
Jattaea sp. bark Zagatala, Azerbaijan [17]
Massarina cf. quercina shoot (phellem) Murg, Switzerland [12]
Mollisia sp. (= Cystodendron sp.) shoot (phellem) Bellinzona, Switzerland [12][41]
Monodictys castaneae shoot (phellem) Bellinzona, Switzerland [12]
Mucor fragilis bark Vinhais, Portugal [24]
Neocucurbitaria cava * leaf or gall Southern Tuscany, Italy [14]
Neopestalotiopsis sp. bark Asturias, Spain [19]
Neopestalotiopsis zimbabwana bark Asturias, Spain [19]
Nigrospora sp. several plant parts Eurobin and Monbulk, Australia [21]
Ophiovalsa cf. suffusa shoot (lenticel) Bellinzona, Switzerland [12]
Paraconiothyrium brasiliense bark Vinhais, Portugal [24]
branch Astroni Nature Reserve, Italy [31]
Penicillium glabrum bark Vinhais, Portugal [24]
Penicillium sp. several plant parts Eurobin and Monbulk, Australia [21]
stem Black Sea region, Turkey [15]
bark Marche, Italy [18]
bud Ankara, Turkey [23]
bark Valpaços, Portugal [24]
branch Astroni Nature Reserve, Italy [31]
Pestalotiopsis sp. leaf Vejoris, Spain [27]
bark Asturias, Spain [19]
bud Ankara, Turkey [23]
Pezicula cinnamomea shoot (phellem) Bellinzona and Murg, Switzerland [12]
Phaeococcus sp. shoot (phellem) Murg, Switzerland [12]
Phoma sp. shoot (phellem) Bellinzona, Switzerland [12]
several plant parts Eurobin and Monbulk, Australia [21]
bud Ankara, Turkey [23]
Pilidiella castaneicola * shoot Bellinzona and Murg, Switzerland [12]
Ramichloridium sp. shoot (phellem) Murg, Switzerland [12]
Rhizoctonia sp. shoot (phellem) Bellinzona, Switzerland [12]
Rhizopus sp. bark Vinhais, Portugal [24]
Sclerotinia pseudotuberosa bark, bud, fruit Viterbo province, Italy [42]
Sordaria rabenhorstii * bark Valpaços, Portugal [24]
Sordaria sp. several plant parts Eurobin and Monbulk, Australia [21]
leaf or gall Southern Tuscany, Italy [14]
Stemphylium vesicarium leaf or gall Southern Tuscany, Italy [14]
Trichoderma atroviride scion Ticino, Switzerland [22]
leaf Vejoris, Spain [27]
Trichoderma hamatum shoot Ticino, Switzerland [22]
Trichoderma koningiopsis bark Shaki, Azerbaijan [17]
Trichoderma sp. stem Black Sea region, Turkey [15]
bark Qakh, Azerbaijan [17]
bark Marche, Italy [18]
Trichothecium roseum leaf or gall Southern Tuscany, Italy [14]
Umbelopsis isabellina bark Balakan and Qabala, Azerbaijan [17]
bark Vinhais, Portugal [24]
Xenoacremonium falcatum bark Balakan and Qabala, Azerbaijan [17]
Xylaria sp. branch Astroni Nature Reserve, Italy [31]
Castanea crenata
Alternaria sp. leaf Kashiwa, Japan [43]
Astrocystis sp. leaf Kashiwa, Japan [43]
Aureobasidium sp. leaf Kashiwa, Japan [43]
Botryosphaeria dothidea leaf Kashiwa, Japan [43]
Colletotrichum acutatum leaf Kashiwa, Japan [43]
Colletotrichum gloeosporioides leaf Kashiwa, Japan [43]
Diaporthe sp. leaf Kashiwa, Japan [43]
Discula sp. leaf Kashiwa, Japan [43]
Glomerella sp. leaf Kashiwa, Japan [43]
Gnomoniopsis castaneae * bark, flower, leaf several locations in New Zealand [13]
Induratia fengyangensis * leaf Kashiwa, Japan [43]
Nigrospora sp. leaf Kashiwa, Japan [43]
Pestalotiopsis sp. leaf Kashiwa, Japan [43]
Phyllosticta capitalensis leaf Kashiwa, Japan [43]
Xylaria sp. leaf Kashiwa, Japan [43]
Castanea dentata
Acremonium implicatum * stem Michigan and Wisconsin, USA [20]
Alternaria alternata stem Michigan and Wisconsin, USA [20]
Alternaria brassicae stem Michigan and Wisconsin, USA [20]
Aspergillus tubingensis stem Michigan, USA [44]
Biscogniauxia aff. mediterranea stem Michigan and North Carolina, USA [44]
Botryosphaeria sp. stem Michigan and Wisconsin, USA [16][20]
Daldinia aff. childiae stem Michigan, USA [44]
Didimostylbe sp. stem Wisconsin, USA [16]
Diplodia corticola stem Michigan and Wisconsin, USA [20]
Diplodia seriata stem Michigan and Wisconsin, USA [20]
Dothiorella sp. stem Wisconsin, USA [16]
Epicoccum nigrum stem Wisconsin, USA [16]
stem Michigan, USA [44]
stem Michigan and Wisconsin, USA [45]
Fusarium sp. stem Massachusetts, USA [46]
Gnomoniopsis castaneae * flower, leaf Ohaupo, New Zealand [13]
stem Michigan and Wisconsin, USA [20]
Mucor circinelloides stem Michigan and Wisconsin, USA [20]
Mucor fragilis stem Michigan and Wisconsin, USA [20]
Nectria cinnabarina stem Michigan and Wisconsin, USA [20]
Nigrospora aff. oryzae stem Michigan, USA [44]
Paraconiothyrium sp. stem Wisconsin, USA [16]
Penicillium glabrum stem Michigan and Wisconsin, USA [16][20]
Penicillium spinulosum stem Michigan and Wisconsin, USA [16][20]
Pestalotia sp. stem Wisconsin, USA [15]
Pestalotiopsis sp. stem North Carolina, USA [44]
Pezicula cinnamomea stem Michigan and Wisconsin, USA [20]
Pezicula ericae stem Michigan and Wisconsin, USA [20]
Pezicula sporulosa stem North Carolina, USA [44]
Strasseria sp. stem Michigan and Wisconsin, USA [20]
Trichoderma atroviride stem Michigan and Wisconsin, USA [16][20]
Trichoderma aureoviride stem Wisconsin, USA [16]
Trichoderma citrinoviride stem Michigan and Wisconsin, USA [20]
Trichoderma harzianum stem Michigan and Wisconsin, USA [20]
Trichoderma sp. stem Massachusetts, USA [46]
Tubakia suttoniana * stem North Carolina, USA [43]
Umbelopsis isabellina stem Michigan and Wisconsin, USA [16][20][45]
Castanea mollissima
Alternaria eichhorniae leaf Qing Long, China [47]
Alternaria sp. leaf Qing Long, China [47]
Auriculibuller fuscus * leaf Qing Long, China [47]
Cercospora canescens twig Qing Long, China [47]
Cercospora sp. twig Qing Long, China [47]
Colacogloea sp. leaf Qing Long, China [47]
Colacogloea terpenoidalis leaf Qing Long, China [47]
Gnomoniopsis castaneae * bark, leaf Ohaupo, New Zealand [13]
Kondoa sorbi leaf Qing Long, China [47]
Kondoa sp. twig Qing Long, China [47]
Papiliotrema sp. leaf Qing Long, China [47]
Phlebia acerina - China [48]
Sporobolomyces roseus twig Qing Long, China [47]
1 Underlined species are also known as disease agents of chestnuts. * Names with an asterisk are the latest accepted for these species, which are different from the ones used in the corresponding references.

This entry is adapted from the peer-reviewed paper 10.3390/plants10030542

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