Dispersal of birch and alder seeds into open areas, number of seeds m-2 year-1
\r\n\tThe book will present up to date knowledge on mentioned ADHD topics in order to be implemented in every day clinical practice.
",isbn:"978-1-83962-495-7",printIsbn:"978-1-83962-475-9",pdfIsbn:"978-1-83962-496-4",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"176f5275d9e1e06b24e0ae07b90c424f",bookSignature:"Prof. Hojka Gregoric Kumperscak",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/9499.jpg",keywords:"Clinical Picture, Symptomatology, Symptoms, Clinical Presentation, Comorbidity, Pharmacotherapy, Nonpharmacological, Nutrition and Diet, Genetics, Neuroimaging, Neurotransmitters, Hormones",numberOfDownloads:528,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"June 10th 2020",dateEndSecondStepPublish:"August 6th 2020",dateEndThirdStepPublish:"October 5th 2020",dateEndFourthStepPublish:"December 24th 2020",dateEndFifthStepPublish:"February 22nd 2021",remainingDaysToSecondStep:"7 months",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:null,kuFlag:!1,biosketch:"Prof. Kumperscak, MD, PhD graduated from the Faculty of Medicine in Ljubljana, Slovenia. She was trained in child and adolescent psychiatry in Slovenia and abroad. She has held the Chair of the Department of Psychiatry in the University of Maribor in Slovenia (2017) and has been Head of the Child and Adolescent Psychiatry Unit, University Clinical Center in Maribor (2008). She is a President of the Slovenian Association for Child and Adolescent Psychiatry and Adolescent Identity Treatment psychotherapist.",coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"53417",title:"Prof.",name:"Hojka",middleName:null,surname:"Gregoric Kumperscak",slug:"hojka-gregoric-kumperscak",fullName:"Hojka Gregoric Kumperscak",profilePictureURL:"https://mts.intechopen.com/storage/users/53417/images/system/53417.jpg",biography:"Prof. Hojka Gregoric Kumperscak, MD, PhD was born in Maribor, Slovenia in 1970. She finished Faculty of Medicine in Ljubljana, Slovenia in 1996. She was trained in child and adolescent psychiatry in Slovenia and abroad (Italy, UK, Germany and Switzerland). \r\nShe has held the Chair of the Department of Psychiatry in the Faculty of Medicine, University of Maribor in Slovenia, since January 2017, and has been Head of the Child and Adolescent Psychiatry Unit, University Clinical Center in Maribor since 2008. She is a President of Slovenian Association for Child and Adolescent Psychiatry and Adolescent Identity Treatment psychotherapist. 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From chapter submission and review, to approval and revision, copyediting and design, until final publication, I work closely with authors and editors to ensure a simple and easy publishing process. I maintain constant and effective communication with authors, editors and reviewers, which allows for a level of personal support that enables contributors to fully commit and concentrate on the chapters they are writing, editing, or reviewing. I assist authors in the preparation of their full chapter submissions and track important deadlines and ensure they are met. I help to coordinate internal processes such as linguistic review, and monitor the technical aspects of the process. As an ASM I am also involved in the acquisition of editors. Whether that be identifying an exceptional author and proposing an editorship collaboration, or contacting researchers who would like the opportunity to work with IntechOpen, I establish and help manage author and editor acquisition and contact."}},relatedBooks:[{type:"book",id:"6550",title:"Cohort Studies in Health Sciences",subtitle:null,isOpenForSubmission:!1,hash:"01df5aba4fff1a84b37a2fdafa809660",slug:"cohort-studies-in-health-sciences",bookSignature:"R. Mauricio Barría",coverURL:"https://cdn.intechopen.com/books/images_new/6550.jpg",editedByType:"Edited by",editors:[{id:"88861",title:"Dr.",name:"R. Mauricio",surname:"Barría",slug:"r.-mauricio-barria",fullName:"R. 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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"878",title:"Phytochemicals",subtitle:"A Global Perspective of Their Role in Nutrition and Health",isOpenForSubmission:!1,hash:"ec77671f63975ef2d16192897deb6835",slug:"phytochemicals-a-global-perspective-of-their-role-in-nutrition-and-health",bookSignature:"Venketeshwer Rao",coverURL:"https://cdn.intechopen.com/books/images_new/878.jpg",editedByType:"Edited by",editors:[{id:"82663",title:"Dr.",name:"Venketeshwer",surname:"Rao",slug:"venketeshwer-rao",fullName:"Venketeshwer Rao"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"4816",title:"Face Recognition",subtitle:null,isOpenForSubmission:!1,hash:"146063b5359146b7718ea86bad47c8eb",slug:"face_recognition",bookSignature:"Kresimir Delac and Mislav Grgic",coverURL:"https://cdn.intechopen.com/books/images_new/4816.jpg",editedByType:"Edited by",editors:[{id:"528",title:"Dr.",name:"Kresimir",surname:"Delac",slug:"kresimir-delac",fullName:"Kresimir Delac"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"19073",title:"Supplying Biomass for Small Scale Energy Production",doi:"10.5772/18581",slug:"supplying-biomass-for-small-scale-energy-production",body:'Our sources of energy are constantly changing. In Sweden the focus is on nuclear and hydro power for producing electricity and total Swedish energy production amounts to about 612 TWh (Anon, 2010). Since Sweden has a cold climate, there is a high demand for energy to heat homes and energy sources other than oil and coal are required. Currently, fuel systems are based on oil and electrical power but there has been an increase in the use of biomass during recent decades. The support of biomass for heating provides 19% of the total Swedish energy output, (Fig. 1).
For centuries trees have been used in a domestic context for firewood and charcoal production. In Sweden, conventional forest management combined with bioenergy production has been practiced for the last 40-50 years. Currently, for economic reasons, bioenergy harvesting is mainly based on large areas of forest land. Tops and branches are harvested from clear cut areas and this biomass contributes greatly to the production of bioenergy. Special equipment is used to harvest biomass, which is used for energy production in direct heating plants. The infrastructure is well established. Most of the harvested material goes to heating plants close to cities, although some is used by individual households.
Total energy use in Sweden in 2007 (Anon, 2010)
The management of forests is mainly directed towards producing pulpwood and timber. The remaining parts of the tree – branches and tops – represent raw material for bioenergy production. Over the last twenty years there has been an increased willingness to make use of these parts of the tree.
Biomass production on former farmland, using willows, poplar and hybrid aspens, is another option for energy production. In general, the Swedish people look favorably on such land use, as well as forest biomass production. There is strict regulation of the management of forest land to minimize the risks of nutrient loss, but no such regulations exist for farmland. Farmers and some sections of the public wish to maintain farmland as an open landscape and to continue with agricultural cultivation.
The Swedish government has twice proposed a reduction in farmland available for the production of cereals, in 1969 and 1986. The plan was to reduce the area by about one million hectares, out of the total of three million hectares. Both attempts failed, although since 1968 350,000 ha have been taken out of production. Some areas of this former farmland have been planted, mostly with Norway spruce and birches, but more than 200,000 hectares which were taken out of production in the period 1970-1980 have received no subsequent management. Today these areas are covered by broadleaved trees with a range of numbers of stems per hectare (Johansson, 1999a), but they are not managed to generate forest products.
Currently, there are standard practices for the management and harvesting of biomass from large forest stands, used in state forests and by forestry companies. It is much more challenging, however, for small-scale forest owners to utilize forest biomass for bioenergy. The amount of biomass that can be harvested from forest land or farmland depends on various factors including site condition, species and management intensity. Few practical recommendations for small-scale owners have been published, and land owners may be unaware of appropriate practice. More information would enhance the use of resources available for bioenergy production.
Herein I present examples of activities and the management of farmland and forest land demonstrating how an owner can undertake small scale biomass production for their own consumption or to supply a local market (neighbors etc.).
The examples presented are:
ingrowth, i.e. natural establishment of broadleaved trees on former farmland via seeds, sprouts or suckers;
direct seeding on farmland;
management of existing mixed stands;
harvesting tops and branches after clear cutting; and
establishing and using fast-growing species.
Finally, some recommendations for small scale bioenergy production are presented.
The most important factors affecting the colonization of open areas by plants are: the year and season of abandonment; the physical state of the site; climate; soil; the existing flora and fauna; proximity and position of source material; opportunities for vegetative regeneration; and the presence, within a range possible for seed dispersal, of an efficient generative reproduction and a rapid, rich and long-distance dispersal of seeds (Falinski, 1980; Harmer et al., 2001). Reviews by Osbornova et al. (1990) and Myster (1993) report many studies of tree generation on abandoned farmland. Natural colonization by trees and other species have been recorded since 1882 at the Broadbalk Wilderness, UK, which has established on former farmland (Harmer et al., 2001). The first tree plants were recorded 30 years after abandonment, i.e. in 1913. The main species regenerating in the area were: common ash (Fraxinus excelsior L.); sycamore (Acer pseudoplatanus L.); field maple (Acer campestre L.); suckers of wild cherry (Prunus avium L.); blackthorn (Prunus spinosa L.); pedunculate oak (Quercus robur L.) and hazel (Corylus avellana L.). In 1998 the dominant and most frequent tree species were pedunculate oak, common ash, wild cherry and sycamore.
Naturally seeded birch (left), sucker from aspen (right) and naturally seeded grey alder (below)
The area of farmland no longer in agricultural production increases as land owners cease activities or direct their energies towards other forms of management. When farmland is abandoned it is invaded by herbs and broadleaved tree species (alder, aspen and birch). In general, one species dominates in the new stand. Most such farmland areas are owned by private individuals. In Sweden, Johansson (1999a) found up to 10,000 broadleaved tree stems ha-1 on about 100,000 hectares of former farmland.
Natural tree establishment in an open area is a slow process, and it may be 5-10 years before trees 2-5 old are seen (Werner and Harbeck, 1982). Most such areas in Northern Europe are small, amounting to 0.5-2.0 ha. In the initial phase, the areas are not noticeable from the surroundings, but later a dense stand is established and the landscape is changed. In general, these areas continue to develop unnoticed by the owner or the public. Eventually, former open areas become covered by forest. Such ingrowth can be the result of natural seeding, sprouting or suckering (Fig. 2).
To produce conditions that will encourage establishment of a wide range of seedlings through natural seeding, and avoid revegetation failing, an understanding of certain abiotic and biotic factors is required. The main factors that affect establishment through natural seeding are: species present, soil type, moisture, competition by grasses and herbs, available seed trees, and weather conditions (heat, dryness etc). It is important to know the timing and periodicity of seed production and dispersal. Basic knowledge about the period for the high rates of seed dispersal is necessary when practicing natural regeneration. In order to encourage natural seeding, ground preparation must be undertaken prior to seed dispersal.
Specific characteristics of a species, such as number of seeds per tree, seed weight and frost resistance, greatly influence the establishment of seedlings. Seeds from some species are wind dispersed (e.g. birch and sallow (Salix caprea L.)) and others water dispersed (e.g. alder); a combination of methods may be used. Studies of wind-mediated seed dispersal for different species indicate the following order of decreasing dispersal:
birch>elm=maple>alder>hornbeam>beech>oak (Augspurger and Franson, 1987; Okubo and Levin, 1989; Willson, 1990; Karlsson, 2001). Table 1 contains data on birch and alder seed dispersal.
Distance from forest stand, m | Country | Reference | |||
<50 | 50-100 | 100-150 | "/>150 | ||
Birch | |||||
"/>400 | "/>100 | Sweden1 | Fries (1982) | ||
"/>200 | <100 | Sweden2 | Björkroth (1973) | ||
58 % of total | 10 % of total | USA3 | Björkbom (1971) | ||
10,450 | 4,200 | 400 | USA4 | Hughes and Fahey (1988) | |
Alder | |||||
78-94 % of total | Sweden5 | Johansson and Lundh (2006) | |||
90 % of total | Sweden5 | Karlsson (2001) |
Dispersal of birch and alder seeds into open areas, number of seeds m-2 year-1
Both downy (Betula pubescens Ehrh.) and silver (Betula pendula Roth) birch produce many seeds. In Estonia, Uri et al. (2007) recorded 3060-36,200 8-year-old birches ha-1 that had been produced by natural seeding on farmland. Seeds from a birch growing at the edge of a clear cut area have been found to spread at a rate of about 100 seeds m-2 up to 200 m from the tree (Fries, 1984). Most of these birch seeds were dispersed during September, although the process continued until December. In a study of sweet birch (Betula lenta L.), Matlack (1989) reported seed were dispersed 3.3 times further than the distance measured by Fries (1984). In a study of silver birch in Estonia, 21 % of the seeds were dispersed in July, 77 % in August and 2 % in September (Kohh, 1936). Heikinheimo (1932, 1937), who reported the same dispersal periods, commented that the weather during summer and autumn is the main factor affecting the period of seed dispersal. Graber and Leak (1992) presented a study on seed fall for broadleaved species in New Hampshire. The mean seed fall (million ha-1) in a study lasting 11 years was: 6.58 for yellow birch (Betula alleghaniensis Britton); 6.38 for paper birch (Betula papyrifera Marsh.); 4.11 for sugar maple (Acer saccharum Marsh.); and 0.17 for American beech (Fagus grandifolia Ehrh.). The seed viability was 30-50 %, depending on species.
Besides wind dispersal, there are some reports of secondary dispersal of seeds (Hesselman, 1934; Matlack, 1989; Greene and Johansson, 1997). The most common is by movement on snow, but for this to occur, seed fall must happen during winter months when snow is on the ground. The seeds can be damaged by friction on frozen snow, thus reducing viability.
The level of seed production by alder depends on the number of hours of sunshine in the period April-September in the year before fruiting, the number of hours of sunshine in the seeding year and the level of seed production in the preceding year (MacVean, 1955). According to MacVean (1955), common alder (Alnus glutinosa (L.) Gaertner) seeds are generally dispersed within a radius of 30-60 m of the mother tree. Karlsson (2001) found that 50 % of the total number of alder seeds produced fell within 5 m and 90 % within 20 m of the stand. In a study by Johansson and Lundh (2006), 50 % of the common alder seeds were found to have fallen before December and 75 % before February. Alder seeds can also be transported by water in spring at the time of snow melt.
Seeds from European aspen (Populus\n\t\t\t\t\ttremula L.) are extremely small (low weight) with a limited growing capacity (Blumenthal, 1942, Latva-Karjanmaa et al., 2006). A large aspen growing close to Tartu city, Estonia, produced 49 kg or 54 million seeds (Reim, 1930). Only a small proportion of the aspen seeds produced will grow; success depends on site conditions, seed size and the level of competition. Aspen seeds can grow on poor sandy sites, burned areas and small patches without vegetation (Blumenthal, 1942). Seeds of sallow are also small and have a plume to aid dispersal (Grime et al., 1988). Seeds of both species can be dispersed over long distances.
The most favorable soil types for rapid establishment of seedlings are fine sand, silt and light clay, sandy-silty till and light clay till. Even peat soils can provide an ideal site, providing there is sufficient water. A mixture of mineral soil and humus is common on farmland, where the area has been cultivated for many years.
Birch seeds establish well on undisturbed sites with a high level of moisture (Mork, 1948; Fries, 1982). During the first part of the growing season in Nordic countries (April-May) soil moisture tends to be low. The lack of rain combined with the sunshine during this period results in a dry soil. Therefore any soil treatment (plowing, harrowing or screefing) should be undertaken in autumn or very early in spring. Studies to determine the best soil treatment to ensure limited cover of competitive vegetation indicate that removal of topsoil is preferable (Karlsson, 1996).
The main difference between sprouting and suckering is that sprouts emerge from a stump whilst suckers originate from roots, (Fig. 3). Both types of regeneration result in fast-growing individual stems. In studies of dormant buds on birch, most have been found close to the ground: 0-10 cm above or 0-5 cm below ground level (Kauppi, 1989; Kauppi et al., 1987; 1988\n\t\t\t\t\tJohansson, 1992a). The number of sprouts per living birch stump has been found to vary between 1 and 52, mean 10±8, decreasing to 3-8 sprouts per stump after five years (Johansson, 1992 b, c). Rydberg (2000) found the number of birch sprouts had decreased by >40 % of the initial number two years after stump creation nine years after cutting, Johansson (2008) found that the initial number of sprouting birch stumps had decreased to 61 and 55 % respectively for downy and silver birch stumps. In a study of downy birch growing in central Finland, the number of sprouts decreased from an average of 9.5 one year after cutting to 5 after three years and 3 after seven years. The sprouting abilities of red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.), sugar maple and yellow poplar (Liriodendron tulipifera L.) growing in West Virginia were studied by Wendel (1974). After ten years the number of sprouts per living stump was 15-20 % of the initial number produced. In another study of yellow poplar, the average number of sprouts recorded six years after cutting was 7.0 per stump (Beck, 1977). Sprouting capacity is highest when a tree is young (Johansson, 1992c). Kauppi et al. (1988) reported the poorest sprouting results from old (40 year) downy birch stumps. Older trees have thicker stem bark, so the buds cannot penetrate the bark and develop into sprouts (Mikola, 1942). Sprouting capacity may depend on carbohydrates in the roots. However, Johansson (1993) found no pronounced peaks in the carbohydrate content in birch roots during the year. Sprouting capacity may also depend on the cutting date. Johansson (1992b) found the highest number of living birch stumps producing sprouts cut in all months but June-October. Etholén (1974) found no effect of cutting time on the sprouting ability of young downy birch stumps.
Sprouts of birch (left) and suckers of aspen (right)
In southeastern New York, Kays and Canham (1991) studied the sprouting ability of four hardwood species: red maple (Acer rubrum L.), gray birch (Betula populifolia Marsh.), white ash (Fraxinus Americana L.) and black cherry (Prunus serotina Ehrh.). They reported that gray birch had the highest mortality (87 %) of stumps after cutting in May but the other species only had mortalities of 10-20 % depending on cutting date. In a study of the suckering capacity of parent trees of American beech, a mean of 41,365 (3,924-89,765) suckers ha-1 was found (Jones and Raynal, 1986).
European aspen and trembling aspen (Populus tremuloides Michx.) are two Populus species with a high capacity for sucker production. The number of suckers after cutting the mother tree differs depending on the cutting date (Johansson, 1993) and on site, stand and management factors (Frey et al., 2003). The age of the mother tree also influences the suckering ability (Brinkman and Roe, 1975). A trembling aspen stand was found to produce 8000 suckers ha-1 after cutting (Tew, 1970). In a study by Alban et al. (1994) of trembling aspen growing in Minnesota, the number of suckers the first year after disturbance was >250,000 per hectare. The number had decreased to 40,000 after five years (Stone and Elioff, 1998). Trembling aspen stands growing on similar soils in Minnesota and British Columbia produced 50,000 suckers ha-1 after five years and the mean sucker height was 2.1 m (Stone and Kabzems, 2002). The root system of an individual aspen is widely spread, with root lengths up to 20 m (Reim, 1930). In a Swedish study, about 70 % of the suckers occurred within 10 m of the parent aspen tree (Bärring, 1988). In a study by Johansson (1993) the content of starch in roots of European aspens fluctuated during the year with the lowest levels in May-July. The same pattern has been reported for trembling aspen by Baker (1925), Zehngraff (1946), Tew (1970) and Brinkman and Roe (1975).The lowest content has been recorded in late May and early June. When aspen is cut in the winter the highest numbers of suckers are produced (Stoeckler and Macon, 1956; Steneker, 1976; Peterson and Peterson, 1992). In other studies (Shier and Zasada, 1973; Fraser et al., 2002) on trembling aspen, no relationships have been identified between carbohydrate content in roots and the number of suckers initiated.
Alder regenerate vegetatively by sprouts or suckers depending on species. In a study of red alder (Alnus rubra Bong.), the number of sprouts per living stump ranged between 5 and 9 (Harrington, 1989). In another study of the same species, the number of sprouts was in the range 9-13 (DeBell and Turpin, 1989). According to Rytter (1996), young grey alders (Alnus incana (L.) Moench) produce sprouts after cutting, but the old trees produce suckers. In a Finnish study, grey alder stumps sprouted within three weeks of cutting (Paukkkonen and Kauppi, 1992). Sucker production by grey alder is the main means of vegetative regeneration when the trees are more than 25-30 years old (Schrötter, 1983). In a study of seasonal variation of carbohydrates in the roots of common and grey alders, levels were found to be highest during September-November (Johansson, 1998). In a study of the influence of felling time on sprout and sucker production by common and grey alder, the carbohydrate content in the roots was found to influence biomass production (Johansson, 2009). The highest number of sprouts from common alder stumps was produced after cutting in August-October (23-24 sprouts stump-1). Ten years later, the number of sprouts had decreased to 1.3-2.3 sprouts stump-1. The average number of sprouts on living grey alder stumps was highest after cutting in March (3.0), August (3.4) and September (3.4), with a reduction to an average of 2.0 after five years. The number of grey alder suckers per m2 was highest, 21.0, after cutting in September with a reduction to 1.5 after five years. The recommendation, therefore, is to cut grey alder in August and September ad common alder in August-October when the largest number of sprouts and suckers will result. In a study on the initial sprouting of 4-year-old red alders, the percentage of sprouting stumps was highest when the alders were cut in January (Harrington, 1984).
In a study of the spouting ability of Eucalyptus in plantations, the number of sprouts per living stump varied, but the highest number was 5-6 sprouts stump-1 (Sims, 1999). The stumps have the capacity to resprout several times, depending on their vigor.
When practicing direct seeding on forest land there are practical recommendations considering among others Norway spruce (Picea abies (L.) Karst.), Scots pine (Pinus sylvestris L.), birch, beech (Fagus sylvatica L.) and oak in relation to the target species. There are, however, few recommendations available for seeding on farmland, although the factors associated with successful establishment are the same as for natural seeding (species, mineral soil, moisture, competition by grasses and herbs, and weather conditions).
The success of establishment of seedlings after direct seeding depends on the nature of the soil treatment and the date of seeding. The critical phase is the emergence of seedlings during the first days or weeks after seeding and the moisture conditions in the treated spots. Generally, precipitation is low in late spring and therefore seeding must be undertaken early in spring.
High quality seeds are expensive and therefore a natural seed source close to the planting site can allow collection from mature seed trees of the appropriate species. Birch and alder are suitable species for producing stands for bioenergy harvest, with subsequent vigorous sprouting or suckering. Depending on seeding method the amount of seeds is 0.5-1.0 kg ha-1.
Using a mixture of species in forest management has been common in Europe for the last three centuries. Hegre and Langhammer (1967) and Stewart et al. (2000) have presented overviews of the importance of mixed stands and their management in different countries worldwide.
Mixed stand of alder and Norway spruce (left), aspen and Norway spruce (middle) and birch and Norway spruce (right)
In Finland and Norway, a forest stand is defined as being mixed if 20 % of its basal area is made up of broadleaved species, with conifers comprising the dominant species (Frivold, 1982). In Sweden, the proportion is 30 % and in Italy 10 % of the basal area. The Swedish definition of a mixed broadleaved and coniferous stand is “a type of stand in which the total percentage of broadleaved species is 30-70 % of the growing stock” (Anon., 2010). In Nordic countries mixed stands are the most frequent type of stand.
Mixed stands mostly establish spontaneously i.e. a planted or naturally regenerated conifer stand is mixed with naturally regenerated broadleaves. Areas of clear felling that are moist are readily colonized by broadleaves, which can establish from seeds, sprouts or suckers. The number of stems can amount to 5000 to 50,000 per hectare. However there is a conflict between broadleaf cover preventing frost damage to young spruce trees and the strong competition between broadleaves and conifer seedlings. In older stands, both species become established, competition is stabilized and the risk of frost damage declines (Johansson, 2003).
Mostly, Nordic forestry is focused on the management of stands for the production of softwood. A large number of young broadleaves are likely to compete with the conifer seedlings in such stands. In the past, the broadleaves were cut or treated with herbicides. Nowadays, with increasing interest in the supply of biomass for bioenergy production, other management systems have been introduced.
When managing mixed forest stands, a stratified mixture of shade-tolerant, late-successional species in the lower stratum and early successional species in the upper stratum is recommended (Assmann, 1970; Kelty, 1992). Mixed stands may contain alder, aspen or birch and Norway spruce (Johansson, 2003), (Fig. 4). The management of mixed stands is often based on stands which have not been cleaned at the correct time. The spontaneous establishment of broadleaved trees takes up to10 years.
A number of methods are practiced in the Nordic countries, most commonly the shelter method (Tham, 1988; Johansson and Lundh, 1991) and the “Kronoberg” method (Anon., 1985). The descriptions in the sections below are based on a mixed stand of birch and Norway spruce, since this is the most common situation, but the same techniques can be used for other broadleaved species with Norway spruce.
When managing this type of stand it is important that the density of the broadleaved stems is not too high once the spruces have been established. According to Braathe (1988), the competition is too strong for spruces if there are more than 1200 birches ha-1 and they are >3 m tall. In that case, he postulated a 30 % decrease in the height increment of the spruce.
This method is common in Finland, Norway and Sweden. It was introduced in Sweden by Tham (1988) with some modifications by Johansson and Lundh (1991). Currently, the same technique is used for birch and Norway spruce in Finland, Norway and Sweden. The principal aim is to create an initial mixed stand with an optimal density of birch.
The method involves two or three steps:
When the spruces are 1.5-2 m tall, the density of birch is reduced by cleaning to 800-1000 stems ha-1.
The “birch shelter” is cut when the birches are 30-35 years old with a diameter at breast height (dbh) of 15-20 cm.
An alternative is to cut all 30-35-year-old birches except 50-100 stems ha-1. The remaining stems should be evenly spread through the stand. These birches will produce high-quality timber during the following 20 years.
This method was first introduced in southern Sweden (Anon., 1985). The aims are to avoid frost damage to Norway spruce plants and to control the number of sprouts that are able to establish after the removal of birch in each step.
The method involves three steps:
When the birches are 3-4 m tall the stand is cleaned. A total of 3000-4000 birch stems ha-1 should be retained. The Norway spruce is not cleaned.
When the birches are 6-9 m tall the stand is cleaned again. A total of 1000-1500 birch stems ha-1 should be retained; the dbh of the birches should be about 5 cm.
When the birch stand is 20-25 years old the birches are felled. They will be 8-12 m tall with a dbh of 8 cm. The mean height of the Norway spruce will be 3-4 m. The spruce stand should be thinned to 2000-2500 stems ha-1.
Alternatively, instead of felling all the birches, 600-800 birches ha-1 could be left for 10-15 years. When the birches are finally cut, their mean dbh will be 15-20 cm.
The most common type of young stands in Nordic countries is mixed birch and Norway spruce, Fig. 5. Many reports describe how to manage birch and Norway spruce. In Finland, Norway and Sweden the management of mixed stands is common (Mielikänen, 1985; Braathe, 1988; Tham, 1988; Mård, 1997; Klang and Ekö, 1999). Frivold and Groven (1996) discussed the importance of managing mixed stands for future high timber quality. The competition between the taller birches and Norway spruce may adversely affect spruce growth. Therefore the birches must be carefully managed with respect to both numbers of stems removed and controlling competition. A common recommendation is to leave 500-1000 stems ha-1 when the birches are 10-15 years old. A Finnish study of a mixed stand of birches (downy and silver) and Norway spruce examined the influence of competition (Valkonen and Valsta, 2001). A reduction of 7-15 % by volume production was reduced by 7-15 % in mixed stands with 1000 birches ha-1 compared to pure spruce stands.
Managed mixed stand of birch and Norway spruce.
Below an experiment in mixed stands of birch and Norway spruce is described (Johansson, 2000b). The experiment was started in 1983 and was based on trials established at eight localities in central and southern Sweden. The experimental stands were 20-30 years old. They were dense, 1520-20,280 stems ha-1, and self regenerated.
The experiment included three thinning regimes:
Thinning of the birch overstory to create a shelter of 500 stems ha-1.
Total removal of the birch trees
Only Norway spruces
At the first cutting, to create the shelter and the pure Norway stands, 1520 to 20,280 birch stems ha-1 with a mean diameter of 5.2 cm were removed. After 5 years, 373 to 507 birch stems ha-1 with a mean diameter of 15.7 cm were recorded.
Data collected five years after the experiment started are presented in Table 2. The competition by the birch shelter did not influence the growth of Norway spruce. As shown in the table, the mean diameter of the Norway spruce trees was almost the same in the shelter as in the pure stands, 7.6 and 7.0 cm respectively.
dbh, cm | Height, m | Stocking level, stems ha-1 | ||
Shelter | ||||
Birch | ||||
Mean ± SE | 13.3±0.4 | 14.2±0.5 | 499±5 | |
Range | 8.1-19.9 | 8.2-20.0 | 480-574 | |
Norway spruce | ||||
Mean ± SE | 7.6±0.3 | 9.7±0.5 | 2811±110 | |
Range | 4.6-9.9 | 5.3-13.5 | 1693-3373 | |
No shelter | ||||
Norway spruce | ||||
Mean ± SE | Mean ± SE | 7.0±0.1 | 8.5±1.0 | 2517±154 |
Range | Range | 3.3-9.2 | 4.2-11.2 | 1293-3453 |
Stand characteristics of the trees remaining five years after cutting
Managed mixed stand of European aspen and Norway spruce
Mixed stands of European aspen and Norway spruce are usually established on rich soils, (Fig. 6). Hegre and Langhammer (1967) and Langhammer (1982) presented results from a Norwegian experiment on farmland that involved planted European aspen and Norway spruce. Aspens and Norway spruces were planted each at a density of 2000 stems ha-1. The aspens were thinned 30 years later and 580 stems ha-1 were retained. Recommendations based on the study stated that planting densities of 2000 Norway spruce and 1000 aspen ha-1 would avoid strong competition by the aspens.
Naturally established mixed stands of alder are common on wet or moist sites, (Fig. 7). Few studies have examined mixed stands of alder and Norway spruce; those which do exist are based on stands that were not managed correctly during the first ten years after establishment (Lines, 1982; Johansson, 1999d).
Managed mixed stand of grey alder and Norway spruce
After clear cutting, tops and branches from felled trees are traditionally left on site together with small trees (Fig. 8). On nutrient-limited sites this slash should not be removed because that would reduce the nutrients present on site. The amount of biomass present in tops and branches is estimated to amount to 20-30 % of the total harvest. The supply of biomass from tops and branches is the main source of bioenergy production in Sweden.
Clear cut area with branches and tops (left) and stacks of branches and tops (right)
Besides conventional forestry management, there is increasing interest in management of so-called fast-growing species. Depending on geographical location, different species can be considered fast-growing. There are at least three types of tree suitable and frequently used for management in Europe, the USA and Canada: Salix clones, poplar and hybrid aspen. In areas with higher temperatures than northern Europe, species of Eucalyptus are also planted.
In Sweden research on short rotations using Salix began in the end of 1900. Today 10,000-15,000 hectares of short rotation Salix stands have been established and are actively managed using advanced technology. The management is based on small-scale plots, where the farmer owns the stand and manages it. Harvesting is undertaken using machinery owned by entrepreneurs and the harvested material is sold to be used for district heating. Common rotation periods are 4-5 years with 5-6 repeated rotations; a plantation lasts a total of 20-30 years before a new one must be established. The plantations must be fertilized and in some cases treated with herbicides. Pathogens (fungi and insects) damaging the leaves and shoots will cause a reduction in growth. As the seedlings represent attractive wildlife habitat, the plantations must be fenced.
Harvested area of Salix (left) and a stack of harvested coppice (right)
Worldwide, and for a long time, poplars have been used for, inter alia, pulpwood and timber production. Currently, short rotation plantations intended for biomass production are being established. In Sweden poplars have been planted in experiments or plots for practical survey for the last 20 years. Poplar plantations covering small areas of 0.5-2 ha on former farmland can produce 80-100 tonnes ha-1 of wood in ten years (Mean annual increment (MAI): 8-10 tonnes ha-1 years-1). If rotations are longer than 10 years, some of the material harvested will be suitable for use as pulpwood. Nowadays short rotation plantations aiming biomass production has been established. In Sweden poplars have been planted in experiments or plots for practical survey the last 20 years. After harvesting, regeneration of older trees by suckers or sprouts is limited. Certain clones and species produce no or only a few sprouts or suckers. This may be because poplars must be young when they are cut for sprouts to be initiated. The bark on the poplar stems is thick already when the alders are 15 years old, preventing any buds from growing into sprouts.
Hybrid poplar stand
Hybrid aspen is a hybrid between European aspen and trembling aspen (Wettstein, 1933). The hybrid was introduced into Sweden in 1939. Today plantations of hybrid aspen are a potential source of bioenergy, pulpwood and timber. The MAI for hybrid aspen is the same as for poplar, 10 tonnes ha-1 year-1. A German study compared the biomass production in repeated five-year rotations of European, trembling and hybrid aspen (Liesebach, et al., 1999). After harvest of the 5-year-old plantation the biomass was: 7 tonnes ha-1 year-1 from European aspen, 18 from trembling aspen and 16-34 from the four clones of hybrid aspen that were examined. The plants were then allowed to produce suckers, resulting in 165,000 suckers ha-1 during the first year and 45,000 suckers ha-1 five years later. During the second rotation, the production was 18 and 20 tonnes ha-1 for European and trembling aspen and 27-41 for the hybrid aspen clones. The amount of biomass after 5 and 10 years could amount to 50 and 100 tonnes ha-1 respectively. If longer rotations are preferred, the focus should be
Hybrid aspen stand
on pulpwood and timber production, with bioenergy derived from tops and branches. After harvesting the trees, the stumps produce 50,000-100,000 suckers ha-1. During the subsequent 5-10 year period the sucker biomass will amount to 50-100 tonnes ha.-1. However biomass production during a 10-year-old rotation was found to amount to 47, 51 and 87-124 tonnes ha-1 respectively for the aspen stands.
The biomass fractions of a tree are the stump (including roots), stem, branches and foliage (needles and leaves). Broadleaved trees and conifers have different fractions of these aboveground components (Johansson 1999a, b). For birches, the mean aboveground fractions are: stem, 75 %; branches, 18 %; and leaves, 7 %. For conifers, the mean values are 63 %, 23 % and 14 % respectively (Johansson, 1999b, c). The percentage represented by needles is higher in young than old conifers, Fig. 12.
Percentage biomass fractions by total d. w. %, of a tree at different diameters (DBH), mm
The effect of repeated harvesting on biomass production and sprouting of downy birches growing in central and northern Finland has been studied by Hytönen and Issakainen (2001). Different harvesting cycles of 1, 2, 4, 8, 12 and 16 years were examined. The main results were that downy birch is not suitable for biomass production using short rotations. Most of the stumps, 87 %, did not sprout in the one year rotations, but 8-year rotations produced the same number of sprouting stumps as the longer rotations.
Reim (1929) reported that European aspen growing along the borders of farmland may produce large numbers of suckers when cultivation ceases. In a study of repeated short rotations of aspen, the number of suckers per hectare decreased with every additional rotation (Perala, 1979). The study included rotations of four or eight years and, in both cases, the number of suckers decreased over the three rotations studied.
There are several establishment and management techniques available that can be applied to small-scale plots for biomass production on farmland and forest land.
The management methods presented here rely on the land owner having extensive and detailed knowledge of biological processes. The changes in growth of individual species and mixed stands must be known. Some of the methods are based on optimal rotation periods and adequate management of the stand, including cleaning and thinning at the correct time. Severe competition could drastically decrease tree growth. Besides the need for the site to be suitable for tree cultivation, the skill of the owners is important. The most important factor, however, is the enthusiasm and curiosity of the owner; without this, most of the methods will not produce the yields suggested in the present study.
Table 3 lists possible future management models for trees established on farmland and forest land. When operating on a small-scale, there are many alternatives and the owner can be more flexible than is possible in large-scale operations. As the possible rotation periods range from 5 to 40 years it is important to have stands of different ages to ensure a continuous supply. Efficient management of such small areas would make it possible to produce a certain amount of biomass for personal use or to sell to neighbors or local heating plants..
Figures for potential energy supply from different stand types and management options allow us to make comparisons and select appropriate ways to use available land.
Most of the methods are cheap, need a short time to establish and involve relatively straightforward management. The raw materials produced can be used to generate energy for the landowner or can be sold.
Activity | Rotation period, years | Biomass, tonnes ha-1 | MWh1 ha-1 | Next generation |
Ingrowth | ||||
Natural seeding | 10-20 | 50-110 | 115-255 | Sprouts or suckers |
Sprouting, suckering | 5-15 | 50-120 | 115-275 | Sprouts or suckers |
Direct seeding | 10-15 | 40-80 | 90-185 | Sprouts or suckers |
Mixed stands | 35-40 | 100-150 | 230-345 | |
Harvesting tops and branches | - | 50 | 135 | |
Fast-growing species | 5-25 | 30-300 | 70-690 | Sprouts or suckers |
Small-scale management of tree stands on farmland and forest land and possible biomass production
The temporal bone is a dense complex bone that constitutes the lower lateral aspect of the skull and has complex anatomy because of the three-dimensional relationships between neurovascular structures. The petrous portion of the temporal bone has a role as the partition between the middle and posterior cranial fossae. It articulates with the occipital bone (occipitomastoid suture) posteriorly, the parietal bone (squamous suture) superiorly, the sphenoid bone (spheno-squamosal suture) and the zygomatic bone (arcus zygomaticus) anteriorly, and the mandible (temporomandibular joint) inferiorly [1, 2]. It contains multiple intrinsic channels, along with the internal carotid artery (ICA), cranial nerves, and sigmoid sinus (SS), all within intricate spatial architecture. Owing to a complex web of foramina and neurovascular structures of the temporal bone, the lateral skull base is a technically difficult region for surgeons. Because the middle and inner ear structures of hearing and equilibrium are preserved in the temporal bone, a surgical dissection of it requires thorough understanding of three-dimensional (3D) map of the topographic anatomy to avoid iatrogenic risks. The relationship between the surface landmarks and expected internal structures and the segmentation of the temporal structures by using key surgical lines and spaces allow a better understanding of its anatomic architecture. Each temporal bone consists of five distinct osseous segments including the squamous, tympanic, petrous, mastoid, and styloid portions [3, 4].
The anterosuperior part of the temporal bone is a large flattened scale-like plate that forms the lateral boundary of the middle cranial fossa. It has three borders and two surfaces [1].
Superiorly, it overlaps the sculpted squamous margin of the middle third of the parietal bone and constructs the squamosal suture. Posteriorly, it forms the occipitomastoid suture with the squamous part of the occipital bone. Also, there is an angle, parietal notch, between the squamous and mastoid portions of the temporal bone (Figure 1). Antero-inferiorly, its thick serrated margin takes part in pterion formation and articulates with the greater wing of the sphenoid bone to form the spheno-squamosal suture. Inferiorly, it fuses and forms the petro-squamous suture with the superior surface of the petrous portion by extending medially as tegmen tympany [5, 6].
The surface landmarks on the squamous portion: 1, temporal fossa; 2, supra-meatal crest; 3, temporal line; 4, external acoustic meatus; 5, supra-meatal triangle (Macewen’s triangle); 6, middle temporal artery; 7, squamo-mastoid suture; 8, mandibular fossa (glenoid fossa); 9, articular eminence; 10, zygomatic process; 11, petrotympanic fissure (Glaserian fissure); 12, mastoid foramen; 13, parietal notch; 14, mastoid process; 15, mastoid notch (digastric fossa); 16, occipital sulcus; 17, tympano-mastoid suture; 18, vaginal process; 19, styloid process.
External surface, the greater part of the temporal fossa, provides origin to the temporalis muscle and is limited below by the curved line, the temporal line, that lies from the supra-meatal crest to the mastoid cortex posteriorly. Below this line, just above and behind the external acoustic meatus (EAM), the supra-meatal triangle (Macewen’s triangle) contains the supra-meatal spine, spine of Henle and the cribriform area (Figure 1). Also, the squamo-mastoid suture is located approximately 1 cm below the temporal line [5, 6, 7]. On this smooth surface, there is a sulcus for the middle temporal artery, which is the medial branch of the superficial temporal artery (STA). Antero-inferiorly, the zygomatic process projects by two roots: the upper border of the posterior root forms the supra-meatal crest and the lower border forms a laterally based projection, known as post-glenoid tubercle or process (PGP). Inferiorly, the concavity along the surface of the anterior root is called the glenoid fossa (GF), which is bounded by the articular eminence (ArE) anteriorly and the PGP posteriorly [5, 6, 7].
Internal surface is rough and concave in shape, and the anterior and posterior divisions of middle meningeal artery (MMA) run in a groove on this surface that defines the boundary of middle cranial fossa with impressions for the gyri of the temporal lobe. Inferiorly, it forms the petro-squamosal suture with the anterior surface of the petrous part [5, 6].
The Macewen’s triangle, a surgical surface marking for the mastoid antrum (MA), is formed between the temporal line superiorly, the posterosuperior wall of the EAM antero-inferiorly, and the opening of the mastoid emissary vein or sinodural angle posteriorly (Figure 1). The temporal line corresponds to the tegmen tympani (TT), which is a bony plate below the middle cranial fossa dura and over the mastoid air cells. The mastoid cortex posterior to the spine of Henle is a guide to the lateral wall of the MA and located 15 mm deep to it in adults but in new born about 2 mm [5, 6, 8]. The cribriform area in Macewen’s triangle is perforated by numerous small holes that serve as a passage for the vessels of the mucosa of the antrum. The dissection along the margins of this triangle is safer because the vital neurovascular structures are absent. Peris-Celda et al. reported that the temporal line is supratentorial and infratentorial in 93% and 7% of the cases, respectively [9]. During retro-auricular mastoidectomy, the MA may be exposured by drilling the cribriform area and provides a safer surgical approach to the tympanic cavity. The tympanic portion and the styloid process may show variations depending on the shape and the position of the spine of Henle. The MA is located in the same line of the spine of Henle at about 10 years; then the MA is enlarged and placed 1 cm behind it [6, 9].
The MMA lies underneath the pterion which is a common junction between the temporal, parietal, frontal, and sphenoid bones. The fracture of this weakest bony part may result in an epidural bleeding. Between the temporal muscle and fascia, the STA and the superficial temporal vein (STV) courses in close proximity with the zygomaticotemporal (ZTN) and the auriculotemporal (ATN) nerves, branches of the trigeminal nerve (TN). Because of a vessel running superficial to the nerve (80% STA), the underlying nerve may be compressed and results in temporal migraine headache. Lee et al. reported that the intersection (compression) point among the ATN, STA, and STV was at an average of 40 mm superior and 10 mm anterior to the tragus, which is a significant surface landmark at the most anterosuperior point of the EAM. The applications of surgical decompression of the ATN in these compression points improve migraine headache [10].
The anterior articular part of the GF is formed by a gentle sloped area of the squamous portion, which facilitates the movement of the temporomandibular joint (TMJ) during wide mouth opening. At the lateral aspect of the ArE, a small bony ridge, articular tubercle (AT), serves as an attachment for the lateral collateral ligament. The PGP inhibits backward displacement of mandibular head and participates to the superior wall of the EAM [8]. The posterior nonarticular part of the GP is formed by the tympanic portion and the squamo-tympanic suture intervenes between them. The inferior edge of the TT (petrous part) divides this suture into two: a petro-squamosal fissure in front and a petrotympanic fissure (Glaserian fissure) behind (Figure 1). The chorda tympani nerve, a branch of the facial nerve, exits the temporal bone through the Glaserian fissure and joins the lingual nerve as the parasympathetic input to start the submandibular and sublingual gland secretions [2, 4, 5].
The articulation between the GF and the condyle of the mandible is called TMJ, which plays an essential role in speech, respiration, swallowing, and specially mastication. Because the TMJ is in close proximity with the MMA, some surgical landmarks around the TMJ and foramen spinosum (FS) play a critical role in surgical approaches. Miller et al. reported that researchers measured the distances from the zygomatic root (first projection of the zygomatic arch = PGP) to some surgical landmarks such as the arcuate eminence (AE), the head of the malleus (HM) under the TT, and the FS to identify the location of the internal auditory meatus (IAM) or the superior semicircular canal (SSC). Also, they described the superior petrosal triangle as a consistent triangle between the zygomatic root, the FR, and the HM to localize the bony tegmen over the tympanic cavity [11]. Baur et al. offered simply identifiable reference landmarks including the AE, the most lateral aspect of the Glaserian fissure and the FS and measured the distances between them to predict the location of the MMA [12]. According to these researchers, the internal landmarks including the HM and Bill’s bar (the vertical crest in the fundus of the internal auditory canal) are in a single plane with the zygomatic root [11].
After the ArE forming the anterior limit of the GF, the anterior root continues in front as a bony ridge that forms the posterior boundary of the infratemporal fossa, which is a small triangular area transmitting the neurovascular structures between the pterygopalatine fossa and temporal fossa. Then, a serrated anterior end of the zygomatic process passes straight forward and articulates with the temporal process of the zygomatic bone and completes the zygomatic arch. The temporal fascia inserts to this arch and the temporal line superiorly and also the masseter muscle origins from the arch inferiorly. The lateral temporomandibular ligament attaches to the AT, and the GF is covered with an articular disc to construct the synovial TMJ with the condyle of the mandible [5, 6, 7].
Anteriorly, the small part of squamous portion takes part in the infratemporal fossa formation with the zygomatic bone and the greater wing of the sphenoid bone. Below the zygomatic bone, the branches of the first and second mandibular parts of the MA with veins and the pterygoid plexus of veins, the mandibular and lingual nerves pass through the infratemporal fossa. During the infratemporal fossa approaching for surgical removal of tumors localized in the orbit, the maxillary and sphenoid sinuses, the detailed anatomical knowledge of these neurovascular structures is needed. Depending on the position of the infratemporal fossa below the floor of the middle cranial fossa and posterior to the maxilla, it is in close proximity with the parapharyngeal and masticator spaces. The parapharyngeal carotid artery enters the carotid canal (CC) behind the FS and foramen ovale. During transpterygoid infratemporal fossa approach, the positions of these surgical landmarks can be used to prevent ICA injury [13].
Ossification of the squamous portion starts intramembranously from one center around the zygomatic process at the 2nd month. At birth it fuses with the other membranous bone, tympanic portion. Normally, at birth the temporal bone consists of three parts; the petrous, squamous, and the tympanic [1].
The mastoid portion forms the pneumatized thick posterior part of the temporal bone. It fuses with the squamous portion antero-superiorly and the tympanic portion anteriorly and the petrous portion anteromedially. It has three borders and two surfaces [5, 6].
Posteriorly, it articulates with the squamous part of the occipital bone between lateral angle and the jugular process and constructs the occipitomastoid suture. Inferiorly, the mastoid process extends as a rough and conical shaped projection and filled with mastoid cells variable in shape and size. Anteriorly, it associates with the tympanic portions of the temporal bone to form the tympano-mastoid suture, and the inferior auricular branch of the vagus nerve (Arnold’s nerve) exits through this suture [5, 14, 15].
Near the squamo-mastoid suture, the occipital belly of occipito-frontalis and auricularis posterior muscles attach on the external surface that is perforated by numerous small foramina. At the posterior border of the mastoid portion or the occipitomastoid suture, the largest one, mastoid foramen is located and transmits an emissary vein connecting the SS with the posterior auricular vein and a branch of occipital artery to the dura mater (Figure 1). The mastoid process serves for the attachment of the sternocleidomastoid, splenius capitis, and longissimus capitis muscles and shows variations in shape and size with respect to sex. The posterior belly of the digastric muscle is originated from the mastoid notch (digastric fossa), which is a depression on the inferomedial margin of the mastoid process (Figure 1). More medial to the notch lies a sulcus, the occipital sulcus, forming a groove for the occipital artery [4, 6].
The internal surface includes a well-defined and curved sigmoid sulcus lying along its junction with the posterior surface of petrous part and lodges the SS, partially the transverse sinus, which are separated from mastoid air cells by a thin plate of bone. The mastoid foramen transmitting the mastoid emissary vein may be open to this sulcus. The SS begins as the continuation of the transverse sinus and lies downward in a S-shaped groove and opens into the superior jugular bulb. There is a sinodural angle between the dura plates of the SS and middle and posterior cranial fossae [2, 5, 9, 16].
The mastoid process shows tree types of pneumatization patterns including pneumatic (full air cell), sclerotic (solid mass of bone), and mixed (air cells and bone marrow) types. Especially, in the anterosuperior part of the mastoid process, there is an irregular cavity that is larger than other mastoid cells and called MA, which corresponds to the cribriform area. It is covered with the mucous membrane of the tympanic cavity and communicates anteriorly with the epitympanic recess of the middle ear via the aditus ad antrum. The tegmen antri, a roof of the MA, separates it from the middle cranial fossa. During embryonic period, the squamous and petrous portions fused each other and forms the petro-squamous suture. In adults, it forms a thin bony septum, the Körner’s septum, by extending into the mastoid process [1, 4, 6, 9, 17]. Körner’s septum divides the mastoid air cells in the mastoid process into a deep petrous part medially and a superficial squamous part laterally. The petro-squamosal sinus or the mastoid emissary vein may infrequently be observed along this septum. During mastoidectomy or transmastoid approaches, awareness of this crucial landmark and its variations is essential to avoid iatrogenic complications. The squamous part starts to develop at 8th week, whereas the petrous part develops later at 6th months during embryogenesis, and each part opens into the MA separately [1]. Also, the mastoid cells are separated by bony plates from the adjacent structures such as the posterior wall of the EAM anteriorly, tegmen plate superiorly, SS posteriorly, digastric ridge inferiorly, and the lateral semicircular canal (LSC) or solid triangle medially. The solid triangle is a compact bony angle between three SCs. During the mastoidectomy, all the air cells around this septum and adjacent bony structures should be removed without damaging the bony plates. To avoid iatrogenic injury to the adjacent structures, the MA must be open superiorly toward TT. The tympano-mastoid suture at the posterior wall of the MA is surface marking of the course of the vertical portion of the facial nerve (FN) [9, 16, 18]. Peris-Celda et al. reported that the parietal notch corresponds to the posterior petrosal point and the SS (the transverse-SS junction) in 66 and 34% of the cases, respectively [9].
Ossification of the mastoid portion is endochondral which is identical to the petrous and styloid portions. At birth, the mastoid process is absent, and the MA is invisible and covered by a thin bony plate that is extension of the squamous portion. At the first year, the mastoid process becomes prominent and the petro-squamous suture arises. The antrum can be seen obviously at about the fifth year. During puberty, the thickness of the process increases, and it becomes pneumatic that is lined by mucous membrane. In adults, the mastoid process may not contain air cells in 20% cases [1, 2, 17].
An annular shaped part of the temporal bone forms the tympano-mastoid suture posteriorly and the squamo-tympanic suture superiorly (Figure 1). Medially, it fuses with the petrous portion, whereas a free lateral part of it constructs the major part of the EAM and also serves an attachment for the cartilaginous part of the external auditory canal (EAC). Its inferior margin is free, and it has two parts on the lateral surface; posterosuperior part forms the EAM, and anteroinferior part limits the mandibular fossa posteriorly [5, 19].
Medially, just above the GF, this suture is subdivided by a thin tegmen part of the petrous portion into two: the petrotympanic fissure posteriorly and the petro-squamosal fissure anteriorly. Lateral part of this upper margin fuses with the back of the PGP to form the nonarticular part of the GF. Inferiorly, the lateral part of the margin gives an attachment for the deep part of the parotid fascia and forms the vaginal process, which wraps the root of the styloid process laterally [2, 4].
Laterally, external surface is bounded by the cartilaginous part of the EAC which extends from the auricle to the tympanic membrane. The EAC is an S-shaped tube, about 2.5 cm in long, that is composed of the lateral third cartilaginous part and the medial two-thirds osseous part [14, 15, 18]. The tympanic part constructs the anterior wall and floor and the lower part of posterior wall of the EAM, whereas the squamous part forms the superior and upper part of the posterior wall of it (Figure 1). The tympanic part grows from the tympanic ring, which is open U-shaped possessing two edge anterior and posterior. The anterior edge forms the tympano-squamous fissure within the anterosuperior part of the EAM and the petrotympanic fissure within the middle ear, whereas the posterior edge forms the tympano-mastoid fissure within the posteroinferior part of the EAM near the stylomastoid foramen (SMF) [2, 4, 19].
The internal surface fuses with the petrous portion and forms the tympanic sulcus for the lodgement of the tympanic membrane, which forms an angle about 55° with the floor of the EAM and separates the external and middle ear (ME). At the upper part, the tympanic sulcus does not fuse each other by forming the greater and lesser tympanic spines and a notch called Rivinus between them. This notch is closed by the pars flaccida of the tympanic membrane. The notch of Rivinus corresponds to the junction between the squamous and tympanic portions [1, 4, 14, 20].
Ossification starts from the four centers around the tympanic ring at the end of the embryonic period (8th week) via intramembranous ossification of the EAM. The tympanic ring at first is nearly straight and then turns into horseshoe shape (annular) and then, the open arms extending upwards terminate in a notch for the location of the tympanic membrane between them. After birth, the upper segment of the tympanic bone grows rapidly but because of the gradual development of the lower segment, a deep notch (tympanic foramen) is left in the anterior part of the bony EAM. Normally, the tympanic ring fuses until the age of 5 year but a dehiscence may persist (range 4.6−22.7%) at the anteroinferior aspect of the EAM, called foramen of Huschke (foramen tympanicum). This fusion defect is not a true foramen, but it may cause a connection between the EAM and the posteromedial part of the TMJ and results in TMJ herniation and the secretion of the parotid gland and also the dissemination of tumor and infections into the EAM [1, 14, 19, 20]. Anteriorly, the EAM may communicate with the retromandibular part of the parotid gland via the fissures of Santorini within the anterior cartilage. Peris-Celda et al. reported that the SSC dehiscence can be observed approximately 1.5 cm posterior to the middle point of the EAM in 86% of the cases [9]. In newborn, the tympanic membrane is infiltrated with air and the tympanic ring forms a bony plate, which may cause the development of a cleft, the auricular fissure, posteriorly and a cleft, the tympano-squamous fissure, anteriorly [19, 20].
The petrous portion is a dense pyramid-shaped bone and composed of the labyrinth of the internal ear, the tympanic cavity of the middle ear and a bony part of the auditory Eustachian tube (ET), and canals for the passage of the ICA and the FN. It is ossified from the otic capsule by forming a 45° angle with the horizontal axis. It has a base, an apex, and three surfaces and three borders [3, 4, 21].
Superiorly, the petrous ridge is the longest border and a boundary between the posterior part of the middle cranial fossa (the anterior surface of the petrous part) and the anterior part of the posterior cranial fossa (the posterior surface of the petrous part). It contains a groove that lodges the superior petrosal sinus (SPS) and the lateral margin of tentorium cerebelli attaches to this margin (Figure 2). Posteriorly, the medial part of the posterior margin articulates with the basilar part of occipital bone along the petro-clival fissure and forms a groove that lodges the inferior petrosal sinus (IPS) that extends from the posteroinferior part of the cavernous sinus to the internal jugular vein (IJV). The lateral part of the posterior margin is free and limits the jugular foramen (JF) supero-laterally and has a triangular notch for the lodgement of the inferior ganglion of the glossopharyngeal nerve (Jacobson’s nerve = GPN). Anterolateral border is formed by the ET extending from the anteroinferior wall of the tympanic cavity to the nasopharynx [3, 4, 9].
The surface landmarks on the anterior surface of the petrous portion: a, petrous ridge (sulcus of the superior petrosal sinus); b, arcuate eminence; c, tegmen tympani; d, sulcus of the lesser petrosal nerve; e, sulcus of the greater petrosal nerve; f, trigeminal impression; g, petrous apex; ıocc, internal opening of carotid canal.
The base is integrated with the inner surface of the squamous and mastoid portions, whereas the apex forms the posterolateral margin of the foramen lacerum (FL) and faces the Meckel’s cave medially. There is a fibrocartilage connection between the apex and the clivus. The internal opening of the carotid canal (IOCC) is observed at the apex for the intracranial entry of the ICA. At the anterolateral part of the FL, the petro-sphenoid ligament connects the tip of the apex to the dorsum sellae of the sphenoid and the abducent nerve lies below this ligament and enters the cavernous sinus adjoining the ICA [1, 7, 16].
Anterior surface describes a triangular area, between the linear lines as follows: a horizontal line that starts from the preauricular burrhole in front of the tragus to petrous apex at the FL and passes through the FS anteriorly, the petrous ridge posteriorly and the petro-squamous suture, which lies along the junction of the petrous pyramid with the vertical part of the squamous portion laterally [3, 16, 22]. It consists of some marking landmarks (Figure 2).
The anteromedial two-third of the musculotubal canal is cartilaginous, whereas the posterolateral third is bony. The bony part consists of two small canals that are separated by a thin bony septum at the lateral part the petrous portion. The tensor tympani muscle passes through the superior semicanal, whereas the inferior semicanal forms the bony portion of the ET. The tensor tympani muscle originates from the greater wing of the sphenoid and inserts into the upper part of the medial surface of the handle of malleus after making a bend around the processus cochleariformis in the tympanic cavity [4, 6]. The ET lies between the tympanic orifice and the isthmus, which has the smallest diameter at the intersection point of the petrous and squamous parts of the temporal bone just behind the sphenoid spine. Brown et al. reported that the ET is subdivided by genu within the membranocartilaginous part into two portions; posterior horizontal ET between the genu and the anterior attachment of the tympanic membrane ridge, whereas the anterior vertical ET lies from the genu to the nasopharyngeal orifice and opens into the nasopharynx. During endoscopic eustachian tube obliteration, the ET is cannulated to treat refractory CSF rhinorrhea by identifying three anatomic parameters: the ET length, isthmus diameter, and genu location. According to a new surgical classification, the cartilaginous portion of the ET is divided into the petrous, lacerum, pterygoid, and nasopharyngeal parts. The bony part attaches to the ET sulcus or sulcus tuba, which is contiguous to the FL medially. The FL is located in the incomplete confluence of the union of the body and the lingular process of the greater sphenoid wing anteriorly, the clivus of the occipital bone medially and the petrous apex posteriorly and covered with the fibrocartilaginous tissue that separates the ET from the ICA [23].
The internal opening of the CC is located near the FL for the passage of the ICA, which is freed at the petrous apex into the cavernous sinus (Figure 2). It is localized medial to the ET, below the greater superficial petrosal nerve (GSPN), a branch of the FN and the trigeminal ganglion [1, 3, 4]. The petrous segment of the ICA within the CC has four anatomic parts, called vertical, posterior genu, horizontal, and anterior genu. During endoscopic endonasal surgery, the junctional part of the ET at the sphenoid spine and FS is crucial landmark to identify and protect the petrous segment of the ICA [13]. The anatomical and surgical relationships between the ET and the petrous segment of the ICA are as follows:
The first curve, posterior genu is located at the level of the bulging basal turn of the cochlea within the bend of the CC. Laterally, the bony part of the ET and the tendon of the tensor tympani muscle; posterolaterally, the promontory and posterosuperiorly, geniculate ganglion are paramount landmarks for the posterior genu of the ICA. The V3 lying anteromedially to the FS and the parapharyngeal segment of the ICA, which passes posteroinferiorly to the sphenoid spine, are critical landmarks. Posterolaterally, the petroclival fissure cartilage is an important landmark to separate the pharyngobasilar fascia from the anterior genu of ICA.
The second turn of the ICA, anterior genu, above the fibrous tissue of the FL is in close proximity to the lacerum segment of the cartilaginous ET laterally and continues as the paraclival ICA in the carotid groove. During the endoscopic approach, the Vidian artery and nerve (VN) are critical landmarks for the second curve of the ICA.
For safe manipulation of the horizontal part of the ICA, the GSPN can be used as surgical landmark. Above the anterolateral margin of the FL the union of the GSPN and the deep petrosal branch of the carotid neural plexus forms the VN which is located anteroinferiorly and lateral to the second turn of the ICA. Malignancies that involve the petrous apex or the carotid artery require the extended endoscopic endonasl approach (EEA). During this procedure, the medial and lateral optico–carotid recesses in the cavernous sinus and the vidian canal (VC) are vital surgical landmarks, which allow to identify the position of the ICA for safe surgical resection near the ICA [13].
At the apex above the CC, a shallow fossa called trigeminal impression (Figure 2) is located for the lodgement of the sensory ganglion of the TN (semilunar ganglion or Gasser’s ganglion) that is covered by a pouch-shaped dura mater called Meckel’s cave [3]. Vascular compression and arachnoid adherence of the TN branches result in trigeminal neuralgia. During endoscopic vascular decompression and Meckel’s cave approaches, the VC, the bone between V2 and the VC and the pneumatization of the sphenoid sinus form a safe route to access and to decompress Gasser’s ganglion with branches, the cranial nerves (III, IV, VI), and the petrous ICA [13, 23].
Behind the trigeminal impression, the roof of the IAM is indicated as a shallow fossa, then it continues with the AE, which is a surgical landmark for the middle fossa approach and located at the junction of the posterior third and the anterior two-thirds of the petrous portion (Figure 2). It is a valuable guide to signify the SSC and the roof of the vestibule up to 93% of the temporal bones [19, 22].
The TT is a thin bony layer covering all of the anterior surface (Figure 2). It forms the roof of the mucosal line including from behind to forward the MA, tympanic cavity and ET which are lined with mucosa. Also, its lateral edge turns downward to subdivide the squamo-tympanic fissure into two parts [1, 3].
On the TT, a bony roof of the geniculate ganglion, there are two foramina, which continue as a small groove adjoining anteromedially; the medial one starts from the hiatus of the facial canal and lodges the GSPN, a branch of the FN and the petrosal branch of the MMA, whereas the lateral one lodges the lesser superficial petrosal nerve, a branch of GPN (Figure 2) [3, 9, 16, 22].
Kaen et al. described the “VELPPHA” area indicating the posterior limit of the transpterygoid EEA. It is composed of the VC (V), the ET (E), the FL (L), the petroclival fissure (P), the pharyngobasilar fascia (PHA), and multiple cartilaginous fibers between them. The posterior opening of the VC, the posterior limit of surgical corridor in the transpterygoid approach, is located above the ET and below the petrous ICA. Behind the posterior margin of the medial pterygoid process, the superomedial border of the ET attaches to the cartilaginous fibers of the FL. The petroclival fissure is situated between the lateral border of the clivus (occipital bone) and the petrous part of the temporal bone and lodges the IPS. The horizontal segment of the petrous ICA turns upward at the medial border of the petrous apex to form the anterior genu of the ICA, and then it continues as the lacerum segment, second vertical segment of the ICA. So, the VC-ET junction is a safe and critical landmark for efficient localization of the lacerum segment of the ICA, as part of the transpterygoid extension of EEA [24].
Tayebi Meybodi et al. described the pterygoclival ligament as a thickened extension of the pharyngobasilar fascia from the pterygoid process to the anteromedial aspect of the lacerum segment of the ICA and reported that the course of the pterygoclival ligament consistently refers to the anteromedial aspect of the lacerum ICA. So, they suggested that the pterygoclival ligament can be used as a safe landmark in case of tumor invasion of the VN, and drilling along the medial aspect of this ligament is more reliable way compared with the VN to avoid the ICA injury during extended EEA. Also, they remarked that this ligament may localize in a venous compartment, which is in contact with the cavernous sinus superiorly and the pterygoid venous plexus posteroinferiorly [25].
The posterior surface, anterior wall of the posterior cranial fossa, is encircled by a venous triangle that is formed by the grooves for SS posteriorly and SPS at the petrous ridge and IPS at the junction of the pars lateralis of the occipital bone and the temporal bone anteroinferiorly. The SS drains into the bulb of the IJV, which exists from the JF together with the cranial nerves (IX-XI) [1, 6, 9].
The IAM is a short canal, about 1 cm long, and has a large orifice, which allows passage of the vestibulocochlear nerve below the FN, the superficial petrosal artery (a branch of the MMA) and the labyrinthine artery (branch of the basilar artery). The bottom (fundus) of the IAM is subdivided into unequal superior and inferior portions by a transverse falciform crest, and into the anterior and posterior portions by a vertical segment, Bill’s bar, respectively (Figure 3) [2, 15]. The localization of the nerves within the IAM is determined by a triangular shaped Bill’s bar as follows; posteriorly the superior and inferior vestibular nerves, anteroinferiorly the cochlear nerve, anterosuperiorly the FN and nervus intermedius pass through the foramina of the fundus (Figure 3) Mortazavi [1, 4, 6].
The aqueductus vestibuli is a bony canal which contains the saccus and ductus endolymphaticus. Its opening is an oblique slit behind the IAM (Figure 3). The endolymphatic sac is located at the lateral part of the posterior surface medial to the posterior SSC [2, 18].
The subarcuate fossa is an indistinct depression (large in new born) located behind the IAM (Figure 3) and transmits a small vein and the subarcuate artery, which is a branch of the meatal segment of the anterior inferior cerebellar artery [4, 5, 9, 14].
The surface landmarks on the posterior surface of the petrous portion: a, petrous ridge; b, arcuate eminence; h, internal acoustic meatus; ı, subarcuate fossa; j, aqueductus vestibuli; k, sigmoid sinus sulcus; m, sulcus of the middle meningeal artery; 12, mastoid foramen.
The inferior surface articulates with the basilar part of occipital bone medially, and the greater wing of the sphenoid bone anteriorly and forms an irregular external surface of the base of the skull. Below the apex, there is a quadrilateral area that serves as an attachment for the levator veli palatini muscle. The lateral part of this area merges with the posterior margin of the greater wing of sphenoid to form the sulcus tuba in front of the cartilaginous portion of the auditory tube [4, 5, 21]. It presents some anatomical landmarks as follows:
The external opening of the CC, which shows an inverted L-shape course, forms the entrance for the ICA, which is surrounded by a plexus of sympathetic nerves (Figure 4). The anterior margin of the horizontal segment of the CC is separated from the musculotubal canal by a thin layer of bone laterally [1, 5, 18].
The jugular fossa is a deep dome-shaped depression at the lateral wall of the JF and located behind the CC and below the floor of the tympanic cavity. It houses the superior bulb of the IJV and the mastoid canaliculus (Figure 4) for the entry of the Arnold’s nerve, which provides sensory innervation of the EAC and auricle [9, 15]. The jugular spine in the jugular notch of the occipital bone divides the JF into the pars nervosa (anterior) and pars venosa (posterior) [4, 5, 9]. Normally, the jugular bulb is located between the IJV and the horizontal course of the SS. Abnormalities of it (80% below the FN in the mastoid cavity) result in dehiscence of the adjacent structures such as: the mediolateral enlargement of the JB results in the vestibular aqueduct, PSC, and IAC dehiscence, whereas the anteroposterior enlargement of the JB may cause the FN dehiscence. Abnormal high riding JB shows both mediolateral and anteroposterior enlargement and results in dehiscence of the FN [26].
Between the jugular fossa and the CC, the inferior ganglion of the GPN is localized in a triangular depression, whereas the inferior tympanic canaliculus penetrates into wedge-shaped bony ridge and transmits the tympanic branch of the GPN and inferior tympanic artery. At the apex of this triangular depression, there is an external opening of the cochlear aqueduct (Figure 4), which connects the perilymphatic space to the subarachnoid space and transmits the cochlear vein [1, 5, 14].
Behind the CC the vaginal process which is the extension of the sharp lower border of the tympanic plate wraps the root of the styloid process (Figure 4). The lower border of that extension serves an attachment for the deep layer of parotid fascia [1, 3, 5, 6].
The surface landmarks on the inferior surface of the petrous portion: FM, fossa mandibularis; FS, foramen stylomastoideum; FJ, fossa jugularis; ET, eustachian tube; eocc, external opening of carotid canal; ıocc, internal opening of carotid canal; star: inferior tympanic canaliculus; arrowhead: cochlear aqueduct.
Internal structures in the petrous portion contain the ME and inner ear. The ME contains an air-filled tympanic cavity and the ossicular chain which is composed of the malleus, incus, and stapes [14]. The walls of the ME:
Lateral wall contains the tympanic membrane and the scutum pointed infero-medially from the squamous portion. The tympanic membrane has two parts; pars flaccida is located in a fibrocartilaginous ring called the tympanic sulcus and susceptible to perforations and pars tensa is situated in the notch of Rivinus above the lateral process of the malleus. At the medial surface of the membrane a depression called umbo is formed by attachment of the manubrium of the malleus.
Medial wall consists of the cochlear promontory, the FC, the oval and round windows. It is divided into three part by the bony ridges: the ponticulus superiorly and the subiculum inferiorly. The oval window (vestibular window) is located above the ponticulus whereas the round window (cochlear window) is below the subiculum, and the tympanic sinus between them is located medial to the FC. The vestibular window is closed by the base of the stapes. The facial recess lies below the lateral SSC and superolateral to the oval window.
Superior wall, the TT, which forms the roof of the ME.
Inferior wall is a bony roof of the IJV.
Anterior wall includes the anterior epitympanic recess superiorly, below it the tensor tympani muscle lies posteriorly and attaches to the neck of the malleus after turning laterally. The orifice of the ET and below it the CC is located inferiorly.
Posterior wall consists of the pyramidal eminence, epitympanum, and facial recess. The stapedius muscle passes through the pyramidal eminence and inserts to the head of the stapes [2, 5, 7, 14, 18].
The tympanic cavity is lined with the mucous membrane that extending into the MA posteriorly and the ET anteriorly. This cavity consists of three parts changing according to the level of the tympanic membrane; the epitympanum (superior to the level of the tympanic membrane), mesotympanum (at the level of the tympanic membrane), and hypotympanum (inferior to the level of the tympanic membrane). The hypotympanum has the orifice of the ET. At the lateral part of the epitympanum below the lateral malleal ligament there is the Prussak space which is bounded by the neck of the malleus medially and the pars flaccida and scutum laterally [2, 3, 5, 14].
Inner ear is comprised of the otic capsule (osseous labyrinth), which surrounds the membranous labyrinth and is divided into three parts from anterior to posterior including the cochlea, vestibule, and three SCs [14]. Cochlea is the spiral shaped bony labyrinth of the inner ear that looks like a snail shell making 2¾ turns about the modiolus and consists of the vestibular and the tympanic and the cochlear ducts, which are formed by an inner membranous partition. The vestibular duct (scala vestibuli) locates at the superior part of the cochlear canal and contains perilymph (rich in sodium ions) and is limited by the oval window, and is separated from the cochlear duct by Reissner’s membrane. The cochlear duct (scala media) locates at the middle part of the cochlear canal and contains endolymph (rich in potassium ions) and is separated from the tympanic duct by the basilar membrane, which has the Organ of Corti including the sensory hair cells. The stereocilia of these cells perceives the potential difference between the perilymph and the endolymph and converts that motion to electrical signals and finally hearing occurs. The tympanic duct (scala tympani) locates at the inferior part of the cochlear canal and contains perilymph as the vestibular duct and is limited by the round window [3, 5, 14, 15]. Vestibule contains the utricle and saccule. SSCs containing three semicircular ducts organized like three flower leafs that join the vestibule. They are located perpendicular to each other; the superior corresponds to the AE, the posterior is parallel to the posterior surface of the pyramid, and the lateral is perpendicular the mucosal plane and angled at 30°from the transverse plane [3, 15].
The FN passes through the anterosuperior part of the IAM and enters the fallopian canal (FC). It contains motor, sensory, and parasympathetic fibers and has six segments as follows:
Cisternal segment lies from the brain stem to the IAM. This part runs together with the cisternal part of vestibulocochlear nerve in same pia mater coverage.
Meatal segment is the smallest part of the FC and contains Bill’s bar as an important landmark.
Petrous (labyrinthine) segment forms first genu (geniculate ganglion) above the cochlea at the lateral wall of the ME and gives a branch named as GSPN. Then, it enters the tympanic cavity and forms an angle ranging from 19 to 107° with tympanic segment of the FC [7, 20]. Because of this segment is the narrowest part and lack of arterial anastomoses, it is susceptible to embolic attacks and vascular compression.
Tympanic segment (first part) starts from first genu and turns backwards to lie in a thin-walled bony canal that runs evenly between the lateral SSC superiorly and the oval window inferiorly and medial to the incus. A dehiscence of the bony canal is more common at this segment in average 41–75%.
Pyramidal segment (second part of the tympanic segment) forms second genu at the posterior wall of the ME above the pyramidal process. It forms an angle ranging from 95 to 125° with mastoid segment of the FC [7, 20].
In the mastoid or vertical segment, the FN gives the acoustic branch for the stapedius muscle, the chorda tympani, and sensitive branch for the auricular region. This segment is located 5.50 mm anteromedially to the SS and extends from the level of the LSC to the digastric ridge (~3.8 mm). Then it exits the temporal bone at the SMF and enters the parotid gland [14, 27].
According to the classical description, the FC has four segments: labyrinthine, tympanic, pyramidal, and mastoid, but the meatal segment is important from an anatomical and surgical perspective. The stylomastoid artery, a branch of the posterior auricular or the occipital arteries, supplies the inferior parts of the FC up to the second genu and anastomoses directly with the petrosal branch of the MMA, which supplies the geniculate ganglion. The FC pathologies are composed of agenesis, aplasia, narrowing, and osteopetrosis of the canal, which result in complete or incomplete facial paralysis. Bell’s palsy depending on the activation of a dormant herpes virus, is responsible for 50% of peripheral FN palsies. The FC dehiscence can be congenital or secondary to the surgical intervention or pathology of adjacent structures and results in cerebrospinal fluid (CSF) otorrhea. Several surgical approaches, including the translabyrinthine, transcochlear and retrosigmoid, are used to treat the FC pathologies [27].
Ossification of the petrous portion begins from the 14 centers that fuse to form otic capsule and is completed at birth. The petrous portion develops from the cartilaginous differentiation of the mesenchyme by endochondral ossification at the 16th week of gestation. The cementum layer in teeth roots and petrous portion of the temporal bone contain the optimal endogenous DNA substrate which can provide information to specify the geographic location for genomic analyses [28]. Damgaard et al. reported that the prevalence of the endogenous DNA contents in nonpetrous bones and teeth is ranged from 0.3 to 20.7%, while the levels for petrous bones ranges between 37.4 and 85.4% [29]. Due to the high density and resistance to harsher climatic conditions of the petrous bone, the otic capsule of the petrous bone preserves DNA substrate extremely well and has much higher endogenous DNA level than the teeth by 5.2-fold on average. So, it is currently acknowledged as the optimal substrates for ancient genomic research [28, 29].
Kawase’s triangle: Borghei-Razavi et al. evaluated the safety of this posteromedial middle fossa triangle for removal of the tumors locating or spreading into the cerebellopontine angle and petroclival area. Kawase’s triangle was identified between the GSPN laterally, the geniculate ganglion at the AE posteriorly, and ganglion gasserian at the trigeminal impression anteriorly. During anterior petrosectomy for accessing the posterior cranial fossa via middle fossa, the GSPN forms the lateral border of the surgical approach (Figure 5) [30].
The surgical triangles on the anterior surface: Kawase’s triangle: Post-med (posteromedial triangle) and Glasscock’s triangle: post-lat (posterolateral triangle). FS, foramen spinosum; GG, geniculate ganglion; TI, trigeminal impression.
Glasscock’s triangle, or the posterolateral middle fossa triangle, is identified between the TN (V3), the geniculate ganglion at the AE and FS (Figure 5). The margins of this triangle are formed by a line between where the GSPN crosses under V3 and the FS medially, a line between the FS and geniculate ganglion laterally, and GSPN describing the base [3, 5, 16].
Rhomboid area (Kawase triangle+postmeatal area) is situated between the GPN, petrous ridge, AE, and the posterior border of the V3. A large tumor located in the midline skull base or spreading into the infratemporal and petroclival region even the cavernous sinus can be removed by extended EEA through V2-V3 corridor to avoid complications including ICA injury, IPS bleeding, TN injury and CSF leak [31].
Trautmann’s triangle is bounded by the SPS superiorly, SS posteriorly, and solid angle which is formed by three SCs anteriorly (Figure 6). In this triangle, the retro-labyrinthine tract from the MA, the endolymphatic sac, and the vestibular aqueduct are located [5, 9].
The surgical triangles on the posterior surface: Trautmann’s triangle margins are formed between the superior petrosal sinus superiorly, the sigmoid sinus posteriorly, and the semicircular canals antero-inferiorly. Star: Citelli’s angle (sinodural angle) is formed between the dural plates of the middle fossa superiorly, the posterior fossa anteriorly and the sigmoid sinus posteriorly.
Donaldson’s line is a surgical line that is parallel to the LSC whereas it is vertical to the posterior SSC and divide it into superior and inferior portions. Below this line medial to the labyrinth the endolymphatic sac is situated. Citelli’s angle (sinodural angle); is bounded by the middle fossa dura plate (SPS) superiorly, posterior fossa dura plate (bony plate covering the MA) anteriorly and the SS posteriorly (Figure 6). During mastoidectomy the air cells in this triangle should be removed [1, 5, 6].
In clinical applications, for fully understanding of the tridimensional architecture of the petrous portion, a reference lines and angles can be defined on the anterior and posterior surfaces from a superior view.
Peris-Celda et al. reported that the EAM and the IAM are located in the same coronal plane on the anterior surface forming surgical triangle [9]. Tawfik-Helika et al. separated the pyramid into four compartments and described two segmentation method to provide better understanding of the distributions of these compartments. They identified four compartments based on their connections: mucosal, cutaneous, neural, and vascular [3, 21].
The mucosal compartment consists of an air filled and mucosa lined cavities from anterior to posterior: the ET, ME, and the MA (Figure 7). The mucosal line in an oblique anteromedial direction extends along these structures and is used for segmental description of this pyramid, and all major anatomical landmarks can be identified relative to this axis for surgical approaches [3, 9, 21].
(A) The margins of the anterior surface of the left petrous portion from a superior view are shown posteriorly by a (thick black) line along the PR, petrous ridge; anteriorly by a (dashed black) line lying from the preauricular burrhole to PA, petrous apex and passing through the FS, foramen spinosum; and laterally by a (dashed white) line along the petro-squamous suture. OC, optic canal; ACP, anterior clinoid process; FL, foramen lacerum; SOF, superior orbital fissure; FR, foramen rotundum; FO, foramen ovale; MMA, middle meningeal artery; IOCC, internal opening of carotid canal; GSPN, greater petrosal nerve; AE, arcuate eminence; TT, tegmen tympani; JF, jugular foramen; IAM, internal acoustic meatus; SSS, sulcus sigmoid sinus. (B) The segmentation of the left petrous pyramid into four compartments including mucosal, cutaneous, neural, and vascular is shown on the left petrous portion.
Extending the mucosal line posteriorly, the MA is separated into medial and lateral parts, whereas anteriorly, the bony portion of the ET is localized at the junction of the petrous and squamous parts and the cartilaginous part opens into the pharynx anteriorly. Medially the line passing through the sulcus of the GSPN and laterally a straight line lying between the foramen ovale and FS are parallel to this line (Figure 7) [3, 9, 21].
The cutaneous compartment is composed of the EAM, which is covered by the skin and separated from the ME by the tympanic membrane medially.
The neural compartment is composed of the otic capsule, which is located medial to ME and the mucosal line. In this bony container, the cochlea, vestibule, and SCs are located from anterior to posterior around the fundus of the IAM (Figure 7).
The vascular compartment is composed of the ICA. The axis passing through the horizontal part of the CC is parallel and medial to the mucosal line (Figure 7) [3]. Moreover, Tawfik-Helika et al. described X and V segmentation methods to advance and enhance education of the compartments.
The X method divides the petrous pyramid into four spaces by using two reference lines intersecting with each other at the ME; the mucosal line and the EAM-IAM line form the X letter (Figures 8 and 9). These four spaces around the ME and the contents in it are as follows:
The anteromedial space—the cochlea and the petrous apex including the ICA
The anterolateral space—the roof of the TMJ
The posterolateral space—the lateral part of the MA
The posteromedial space—the posterior labyrinth and the medial part of the MA
Schematic representations of the segmentation of the left petrous portion by using X and V methods.
Schematic representation of the external and internal landmarks on the left petrous portion. V, trigeminal nerve and branches (V1, V2, V3); TI, trigeminal impression; IOCC, internal opening of carotid canal; ET, Eustachian tube; GG, geniculate ganglion; ME, middle ear; MA; mastoid antrum; EAM, external acoustic meatus; TMJ, temporomandibular joint; SCCs, semicircular canals; IAM, internal acoustic meatus; VII, facial nerve; VIII, vestibulocochlear nerve; IX, glossopharyngeal nerve; X, vagus nerve; XI, accessory nerve.
The V method arranges five segments around the mucosal line (Figures 8 and 9) These five segments and the contents in it are as follows:
The petrous apex segment—the ICA medial to the ET
The otic capsule segment—the IAM, cochlea, vestibule and SCs
The mastoid segment—the angle around the MA
The EAM segment—the lateral part of the ME
The TMJ segment—the roof of the TMJ lateral to the ET [3].
Detailed description of the temporal anatomy pointing to relationships between internal and external landmarks and a holistic approach including X an V segmentation methods that break down the petrous pyramid into spaces and compartments can provide an easy way to understand and to use surgical applications. The compartmental approach can be helpful in the fields of education and radiological applications as well as surgery.
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