Dimensions of the designed polarization diversity antenna.
\\n\\n
Dr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\\n\\nSeeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\\n\\nOver these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\\n\\nWe are excited about the present, and we look forward to sharing many more successes in the future.
\\n\\nThank you all for being part of the journey. 5,000 times thank you!
\\n\\nNow with 5,000 titles available Open Access, which one will you read next?
\\n\\nRead, share and download for free: https://www.intechopen.com/books
\\n\\n\\n\\n
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'
Preparation of Space Experiments edited by international leading expert Dr. Vladimir Pletser, Director of Space Training Operations at Blue Abyss is the 5,000th Open Access book published by IntechOpen and our milestone publication!
\n\n"This book presents some of the current trends in space microgravity research. The eleven chapters introduce various facets of space research in physical sciences, human physiology and technology developed using the microgravity environment not only to improve our fundamental understanding in these domains but also to adapt this new knowledge for application on earth." says the editor. Listen what else Dr. Pletser has to say...
\n\n\n\nDr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\n\nSeeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\n\nOver these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\n\nWe are excited about the present, and we look forward to sharing many more successes in the future.
\n\nThank you all for being part of the journey. 5,000 times thank you!
\n\nNow with 5,000 titles available Open Access, which one will you read next?
\n\nRead, share and download for free: https://www.intechopen.com/books
\n\n\n\n
\n'}],latestNews:[{slug:"stanford-university-identifies-top-2-scientists-over-1-000-are-intechopen-authors-and-editors-20210122",title:"Stanford University Identifies Top 2% Scientists, Over 1,000 are IntechOpen Authors and Editors"},{slug:"intechopen-authors-included-in-the-highly-cited-researchers-list-for-2020-20210121",title:"IntechOpen Authors Included in the Highly Cited Researchers List for 2020"},{slug:"intechopen-maintains-position-as-the-world-s-largest-oa-book-publisher-20201218",title:"IntechOpen Maintains Position as the World’s Largest OA Book Publisher"},{slug:"all-intechopen-books-available-on-perlego-20201215",title:"All IntechOpen Books Available on Perlego"},{slug:"oiv-awards-recognizes-intechopen-s-editors-20201127",title:"OIV Awards Recognizes IntechOpen's Editors"},{slug:"intechopen-joins-crossref-s-initiative-for-open-abstracts-i4oa-to-boost-the-discovery-of-research-20201005",title:"IntechOpen joins Crossref's Initiative for Open Abstracts (I4OA) to Boost the Discovery of Research"},{slug:"intechopen-hits-milestone-5-000-open-access-books-published-20200908",title:"IntechOpen hits milestone: 5,000 Open Access books published!"},{slug:"intechopen-books-hosted-on-the-mathworks-book-program-20200819",title:"IntechOpen Books Hosted on the MathWorks Book Program"}]},book:{item:{type:"book",id:"901",leadTitle:null,fullTitle:"Transgenic Plants - Advances and Limitations",title:"Transgenic Plants",subtitle:"Advances and Limitations",reviewType:"peer-reviewed",abstract:"Development of efficient transformation protocols is becoming a complementary strategy to conventional breeding techniques for the improvement of crops. 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She received her BSc in İstanbul University in 1993, and MSc and PhD from Gebze Institute of Technology in 2000 and 2005, respectively. She carried out her Post-Doc study on plant cryopreservation at CNR, IVALSA, Florence, Italy. She has published more than 75 scientific papers, conference proceedings and book chapters in the field of plant biotechnology including molecular markers, micropropagation, medium and long-term conservation, transgenic research, and somaclonal variation assessment. She is the author of the book entitled ‘Transgenic Plants: Advances and Limitations’. 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\r\n\t“Gastritis” is the more discussed and ambiguous diagnostic definition of the gastric clinicopathological conditions. There are no connections between symptoms, endoscopic and histological features. The clinical and instrumental exams provide data that are difficult to correlate with each other. The symptoms referred to esophagogastroduodenal tract are the epigastric pain, heartburn, dyspepsia; the endoscopy can identify various hemorrhagic and erosive mucosal lesions, hypertrophy, etc. with the subsequent histobioptic ascertainment. Moreover, poor correlations between the endoscopic abnormalities and histological features of the same lesions: gastric mucosa endoscopically as normal should show histologically severe signs of inflammation; the evident endoscopic damage by drugs might turn out as mild phlogistic lesions with histology.
\r\n\r\n\tThe term gastritis should be used in case of histological phlogistic characteristics. However, the endoscopy has a fundamental role in the gastritis diagnosis, because the microscopic evaluation is made on the mucosal biopsies of abnormalities.
\r\n\r\n\tThe classifications of gastritis are based on histology; unfortunately, also within the histological field, it’s difficult to present a classification of gastritis, because there are various criteria: first, histological features of inflammation subdivide acute and chronic gastritis; more detailed histological characteristics do list gastritis due to drugs, chemicals, infection agents, trauma, foreign body, tumors, autoimmune gastritis, vascular gastropathy, granulomatous gastritis, etc.; finally others histological characteristics of development of inflammation differentiate chronic atrophic and hypertrophic gastritis. Finally, the gastritis is an inflammatory lesion of the gastric mucosa with various and numerous etiologic factors and histological features of inflammation; consequently, the gastritis is a pathological state, but not a defined disease.
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He was previously Assistant Professor (1974–1982)\nand Associate Professor (1982–2001) at the School of Medicine and Surgery, University of Bari, Italy. He graduated in Medicine and Surgery(1970) and completed postgraduate training in General Surgery (1975)\nand Emergency Surgery (1979) at University of Bari, Italy. He received\na diploma of 'Maitrise Universitaire en Pedagogie des Sciences de la\nSantè” from the University Paris-Nord Bobigny (1995). His main research\ninterest is hepatobiliarypancreatic surgery, specifically the management\nof acute pancreatitis and treatment of pancreatic and liver tumors. He\nhas published research papers, reviews, congress proceedings, and book\nchapters. He attended, in the period 1991–2016, for short periods every\nyear, the Hepatobiliarypancreatic Surgery Service of Beaujon Hospital,\nUniversitè de Paris, Clichy. He developed a seminar on 'Cystic Tumours\nof the Pancreas” for the Erasmus Program at Ghent University, Belgium, in2010–2011. 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After this allocation, ultra-wideband has received attention from wireless communication experts owing to its advantageous features like wider bandwidth, low cost, low susceptibility to multipath fading, reduced probability of detection and intercept and potentially high data rates. In a highly dense and dynamic environment, the UWB systems suffer from multipath fading due to reflection and diffraction. This multipath fading results into the degradation of signal-to-noise ratio (SNR) and channel capacity.
An effective method to resolve these multipath fading issues is the incorporation of antenna diversity techniques in wireless communication systems. Several types of diversity, such as space/spatial, pattern and polarization diversity, have been already proposed and implemented to receive multiple signals [2, 3, 4].
In a diversity scheme, the power or signal-to-noise ratio of the received signal is optimized by the selection or combining of output signals in several ways like selection combining, equal gain combining or maximal ratio combining. The detailed description of diversity combining techniques is available in [5, 6].
For good diversity performance, the received signals should have very low correlation between them [7]. An increase in correlation reduces the combining efficiency. In a spatial diversity scheme, a large separation (compared to wavelength) between the antennas is used to achieve decoupling between signals. This large space requirement limits the use of this diversity method. To overcome this drawback, other techniques such as pattern or polarization diversity [8, 9] are investigated. These alternate techniques involve the use of two or more antenna elements with different radiation patterns [10]. An UWB system with polarization diversity technique has potential applications in advanced instruments used for microwave imaging, radar and high-speed data transfer. Some UWB polarization diversity antennas are already reported in the literature [11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28]. However, the application of those available structures is limited due to their large dimensions, multilayer structure, complex feedline, complex geometries, etc.
Among the various bandwidth enhancement techniques, the use of fractal geometries is proven to be a good method. Fractal antenna structures have a compact size and wideband performance due to properties of self-similarity, space filling and effective energy coupling properties [29].
In this chapter, a compact CPW-fed UWB fractal antenna with polarization diversity performance is presented. The bandwidth of the antenna structure [29] is enhanced by loading the coplanar ground planes with a quarter wavelength long rectangular notches. Two identical copies of this antenna structure are arranged orthogonally to achieve good interport isolation and orthogonal polarization diversity performance without affecting the UWB performance. In the following sections, antenna design description is followed by discussion of frequency domain analysis results, time domain analysis results and diversity performance parameter calculation. Finally, it is concluded with major findings of this chapter.
The geometry of the antenna structure is demonstrated in Figure 1, and its optimized dimensions are listed in Table 1. It is etched on a 1.6 mm thick FR-4 epoxy substrate having a relative permittivity of 4.4 and loss tangent of 0.02. All the dimensions are optimized by using finite element method (FEM)-based Ansoft’s high-frequency structure simulator (HFSS) [30]. During simulation, the radiating patch and ground planes are assumed to be perfect electrical conductors. The antenna structure is designed in two steps.
Geometry of the antenna.
Dimensions of the designed polarization diversity antenna.
The fourth iterative fractal geometry of radiator is derived from a rectangular monopole by loading it with a pair of triangular notches on its edges. The flow chart of designing the intermediate design steps is presented in Figure 2. The intermediate design steps for radiator geometry are shown in Figure 3. The iteration structure dimensions are governed by Eq. (1):
Flow chart of designing initial radiator geometry.
Designing stages of proposed MIMO antenna element (a) Zeroth iteration, (b) First Iteration (c) Second Iteration (d) Third Iteration (e) Third iteration with ground notch.
where:
n = iteration number = 2 or 3.
R1 = dimension of the first iteration, i.e. L1, W1 and Wm1.
Rn = dimension of the nth iteration.
r = iterative ratio = 0.4.
In the first step, the dimensions of radiating patch are unaltered. The bandwidth of inner tapered tree-shaped fractal antenna is enhanced by loading the ground plane with quarter wavelength rectangular notches to excite an additional resonance at 18.6 GHz. The dimensions of notch are governed by Eq. (2):
Lr = electrical length of slot for resonance.
fr = resonance frequency = 18.6 GHz.
In the second step, two identical structures designed in the first step are placed orthogonal to each other. The air gap between the two structures and their locations are optimized. All the frequency domain results are calculated by using HFSS. The time domain results and diversity performance parameters are analysed by using CST MWS [31].
The intermediate antenna design steps are compared in terms of their S11 curves in Figure 4. It is observed that the bandwidth is increasing with an increase in iteration. For further increase in iteration, no significant improvement is observed. The reflection coefficient curves for the initial antenna structure with and without ground notches are illustrated in Figure 5. Its quantitative analysis is listed in Table 2. It is observed that the lower band edge frequency is negligibly changed, whereas the higher band edge frequency is shifted from 16.4 GHz to 19.4 GHz in the case of notch-loaded ground plane. The initial resonances are slightly shifted to higher frequency with an additional resonance.
Reflection coefficient of intermediate antenna design steps.
Reflection coefficient versus frequency characteristic of single antenna element.
The designed diversity antenna structure is simulated by using HFSS and CST MWS simulators. The variations of simulated scattering parameters with frequency are demonstrated in Figure 6. The quantitative analyses of bandwidth for two antenna elements used in designed antenna structure are presented in Table 3. From Figure 6 and Table 3, it is observed that there are some discrepancies among the two simulation results. These discrepancies can be attributed to the different mesh size suitable for numerical techniques on which the simulators are designed. In addition to mesh size, it is also important to mention that in CST MWS the structure can be solved in single pass instead of solution for different frequency spectrum, i.e. 1–2, 2–4, 4–8, 8–16 and 16–32 GHz in HFSS. The differences between the S11 and S22 characteristics are due to asymmetrical structure with respect to substrate. A good isolation of more than 15 dB is achieved. The designed antenna has resonances at the frequencies of 6, 8, 10.8, 15.8 and 18.8 GHz.
Scattering parameters versus frequency characteristics of the designed antenna.
Bandwidth comparison of single antenna element with and without ground notch.
Bandwidth of two ports.
The comparison among the designed antenna and previously reported polarization/pattern diversity antenna structures is listed in Table 4. It is observed that the designed antenna has wider bandwidth, good isolation and miniaturized dimensions than other structures.
Comparison of designed antenna with other UWB diversity antenna.
The simulated radiation patterns of two antenna elements at the resonance frequencies of 6, 8, 10.8, 15.8 and 18.8 GHz in all three planes are illustrated in Figure 7. From Figure 7, it is observed that the antenna structures have bidirectional and omnidirectional patterns at lower frequencies. At higher frequencies, the patterns are distorted omnidirectional in nature due to the excitation of higher-order modes at those frequencies. It is also clearly observable that the patterns of both antenna structures have a phase difference of 90° in each plane as desirable.
Radiation patterns of the two antenna elements in XY, YZ and ZX planes.
Figure 8 illustrates that the peak realized gain of Antenna I is varying from 0.52 to 4.98 dB with an average of 3.5 dB over the operating frequency band. It also presents that the gain of Antenna II is varying between 1.43 and 3.5 dB with an average of 2.45 dB.
Peak realized gain versus frequency characteristics.
The variations of radiation and total efficiencies with frequency for both antenna elements are shown in Figure 9. The radiation efficiencies of both antenna elements are more than 0.7 with an average of more than 0.86. Similarly, the total efficiencies have an average of 0.83. The efficiencies are decreasing with an increase in frequency due to the use of lossy FR-4 substrate.
Radiation and total efficiencies of the two antenna elements.
During time domain analysis, two identical antenna structures are arranged in two major configurations, i.e. face to face and side by side as shown in Figure 10. In each of these configurations, one port is excited, whereas the other port is terminated with a matched load. The antenna structure is excited with a Gaussian pulse having a centre frequency of 13 GHz and bandwidth of 1–25 GHz. The normalized amplitudes of the transmitted and received signals are presented in Figure 11. From these normalized amplitudes, the correlation between the two signals, i.e. system fidelity factor, is calculated by using Eq. (3). The calculated values of system fidelity factor for four cases are listed in Table 5. The values listed in Table 5 indicate that the signal is slightly distorted in side-by-side configuration in comparison to face-to-face configuration for both cases.
Time domain analysis configurations of the diversity antenna. (a) Face to face (b) Side by side.
Normalized amplitudes of the transmitted and received pulses in all four configurations. (a) Face to face (b) Side by side.
System fidelity factor for four configurations of the designed antenna.
where st(t) and sr(t) are the transmitted and received pulses and τ is the group delay. The variations of group delay with respect to frequency for all four cases are illustrated in Figure 12. It is observed that the group delay has its variations less than 1 ns over the entire band of operation.
Group delay versus frequency characteristics of the diversity antenna for all four configurations.
To analyse the diversity performance of the designed antenna, various parameters like envelope correlation coefficient, diversity gain (DG) and mean effective gain (MEG) are to be calculated from s-parameters or farfield patterns. The envelope correlation coefficient (ECC) signifies the correlation between the radiation patterns of two antenna elements. For the designed antenna structure, ECC (
Envelope correlation coefficient versus frequency characteristic.
From Figure 13, it is observed that the ECC values are less than 0.005 in the entire band of operation. These low values of ECC (<0.5) signify that the designed antenna is a good candidate for the UWB applications with polarization diversity [16].
The mean effective gain measures the antenna gain of each antenna element taking the radiation power pattern effects, the antenna total efficiency and the propagation effects into account. It is calculated by using Eq. (5) and is plotted for each antenna in Figure 14.
Mean effective gain versus frequency characteristics of the designed antenna.
where XPR represents the cross-polarization ratio,
Another important parameter used to identify the suitability of an antenna for diversity applications is diversity gain. It is the difference between the selection combined cumulative distribution function (CDF) and one of the other CDFs at a certain CDF level. The commonly used CDF level is 1% [33]. The DG of the diversity antenna can be calculated approximately by Eq. (6) [34]. From Figure 15, it is observed that the diversity gain value is almost constant in the entire band of operation.
Diversity gain versus frequency characteristic.
In the case of a rich multipath environment, the maximum rate of transmission for reliable transmission in a communication channel is estimated by calculating capacity loss (b/s/Hz). For a MIMO antenna, a channel capacity loss of less than 0.4 b/s/Hz is acceptable [35]. It is calculated by using the correlation matrix (7) [35].
where
i, j = 1 or 2.
Figure 16 shows that the channel capacity loss changes with the variation of frequencies. It can be seen that the capacity loss values are always less than 0.3 b/s/Hz in the UWB operating range.
Capacity loss versus frequency characteristic.
A compact third iteration fractal antenna with notch-loaded ground plane for UWB applications is investigated and analysed. By using orthogonal arrangement of two antenna elements, polarization diversity performance is achieved. The designed antenna has an impedance bandwidth of 4.7–19.4 GHz, isolation of more than 15 dB, a size miniaturization up to 92% over previously reported structures and good diversity performance parameters values like ECC < 0.005, almost constant DG = 10 and channel capacity loss of less than 0.3 b/s/Hz which makes it suitable for UWB polarization applications in future wireless communication systems to mitigate the multipath fading.
Over a large area of the Amazon rainforest, extending from the Brazilian state of Para to the south-east of Peru (Madre de Dios department), passing through Surinam, one can observe curious clay buildings, having the shape of turrets, or chimneys (Figure 1a), with a height of 20 to 40 cm and an internal diameter at their base of about 2 cm (Figure 1b). The turret surmounts a vertical well (Figure 1c) with a depth of up to about a meter, i.e. the thickness of the fertile soil layer. The surface inside the turret is perfectly smooth (Figure 1d).
(a) View of a turret. (b) Turret removed. (c) Entrance to the well. (d) Inner surface of the summit.
Each turret is the visible part of the pupal burrow of the cicada Guyalna chlorogena (Figure 2), or Fidicina chlorogena, according to its old taxon [1]. Endoscopic exploration made it possible to observe the nymph in the well (Figure 3) and to verify that each burrow is occupied by a single nymph, male or female [2], which builds its turret (between December and February) a few months before moulting into a winged imago, and reproducing (between late July and early September).
The cicada Guyalna chlorogena; young imago after moulting (photography Vanessa Gama).
Endoscopic images of the nymph in its well; lbi: Labium, frl: Foreleg,mdl: Middle leg, hnl: Hind leg. Clay can be seen on the head of the nymph.
Our research was conducted at the Museu da Amazônia (MUSA)1, installed in the Botanical Garden of Manaus, on the edge of the Adolfo Ducke Reserve (Figure 4), in the northern area of Manaus, State of Amazonas, Brazil, in an area of about twenty hectares, around 59° 56′ 21.5” WO and 3° 0′ 19.7” S. A total of more than 250 burrows were observed, between November 2013 and September 2019. About 60 buildings were monitored daily. The observations made up to 2016 have been published [2]. The results obtained subsequently will be the subject of a second publication (Béguin, Gama and Ribamar Mesquita Ferreira, to be published).
Manaus and the Adolfo Ducke Reserve. Google Maps https://www.google.ch/maps/place/Reserva+Florestal+Adolpho+Ducke/@-2.8580657,-60.0242213,71,452 m/data =!3 m1!1e3!4 m5!3 m4!1s0x926c1ec4d40d48b7:0x897d42e519777eb2!8 m2!3d-2.9633439!4d-59.9228331.
The turret is constructed from the well, which has a double curvature near the ground surface (Figure 5a) and a single one in depth (Figure 5b), as revealed by cement casts.
(a) concrete moulding inside an 88 cm deep well, 26 cm of which have been excavated. The curvature (circle), just below the ground level, is clearly visible. The turret is traced (tur). (b) oblique galleries (gal) at the base of the well, oblique as well, up to a curvature at 15 cm from the bottom; cem: Cement moulding.
From the moment it appears, the night-time growth of the turret is very rapid; 3–4 cm per night. The nymph uses a special technique that allows it to lengthen the top without ever opening it, so without exposing itself to predators. It softens the top with a mixture of its urine and clay drawn from the bottom of the well and loaded on its clypeus; then it pushes everything upwards [2].
When the nymph encounters obstacles, it continues its construction obliquely, but restores the verticality as soon as possible, manifesting an acute perception of gravity (Figure 6).
A pile of stems and leaves became an obstacle during the construction of this turret; the nymph avoided it by raising the turret obliquely, but it reestablished the verticality as soon as the obstacle was passed.
The monitoring of many buildings, after the construction of the turret and until the nymph abandons its burrow before moulting into a winged imago, has made it possible to identify various maintenance and rehabilitation behaviours, as well as to make hypothesis about the role of the turret.
If a turret is damaged, the nymph fixes it without delay. It performs an occlusion with a mixture of clay and urine (Figure 7) if its turret was severed, before restoring the initial height (see below). Maintaining the sealing of the building appears to be a priority for the nymph; if one experimentally fractures a turret and then re-stack pieces over the base, the nymph plugs the interstices (Figure 8) by injecting soggy clay with its urine.
Occlusion in progress by the nymph with clay mixed with its urine (mcl) after its turret has been severed.
Sealing of interstices by injection of soaked clay (cl).
When a turret has been destroyed, or even when it tips over (Figure 9), the nymph rebuilds it completely.
Reconstruction of the turret after failover.
The experiment was carried out [2] to section a turret experimentally and to continuously monitor the repair; the nymph clogs the section and restores the original height within days using the same lengthening technique as when growing, and at the same rate of about 3 cm per night. The result of this experiment gives credence to the idea of the requirement of a minimum height necessary to maintain appropriate parameters (humidity, pressure, and O2 and CO2 levels) for the survival of the nymph in its burrow.
A spontaneous increase in turret height to a value held constant thereafter has occasionally been observed, reflecting an increase in the minimum required height discussed above.
A spontaneous decrease in the height of the turret can also be observed, resulting from building a summit inside the turret, below the existing one, which, probably because it is no longer in contact with the moisture inside the burrow, dries up and crumbles (Figure 10), revealing the new top. After a few days, the parts above the new summit have completely disappeared and the result is a turret with reduced height.
Spontaneous reduction in the height of the turret by construction of an internal top and drying out of the old one. The white line represents the old wall of the upper part of the turret.
The reasons leading to the performance of this operation by the nymph remain to be elucidated because, during experiments of artificial lengthening of a turret by transplanting fragments from another one, a nymph does not manifest a requirement of a maximum height; in the long run, it accommodates a higher turret than the one it has built itself; it does not practice the technique which has just been described, nor any other. On the other hand, it ensures the sealing of the modified turret by plugging with soaked clay the interstices between the base of its turret and the implants received, in the same way as described above (Figure 8).
This action of the nymph is rarely observed, but its peculiarity makes it worthy of presentation. A turret emerges near another which later tips over and lies down on the ground. The communication with the well is either completely closed or in the process of being sealed with clay (Figure 11a). The hypothesis can be put forward (Figure 11b) that the new turret is built over a deviation made by the nymph from the exit of its well.
(a) Spontaneous reconstruction of a new turret. (b) Diagram of deviation by the nymph of the outlet of the well: obt: Filling with clay of the old outlet, dfl: Deviation made for a new outlet, ft: Former turret, nt: New turret.
A correlation has been established [2] and confirmed later (Béguin, Gama and Ribamar Mesquita Ferreira, to be published) between the appearance of intense rains and the simultaneous temporary opening of the turrets at their top. A succession of 3 episodes, between July 19 and 23, 2016, turned out to be particularly significant; many openings (Figure 12) appeared, which were then closed when the precipitation stopped (Figure 13). The openings vary in shape, from a small hole about 5 mm in diameter (Figure 12a) to a larger opening where the contour of the clypeus, eyes or anterior part of the pronotum (Figure 12b) can frequently be observed, as a consequence of the liquefaction of the clay from the top wall by applying these parts covered with clay wet with urine. After the rain has stopped, the nymph closes these openings by injecting moist clay (Figure 12c–f).
(a) Circular orifice resulting from a slight pressure of the mass of soggy clay against the wall of the summit. (b) Orifice with imprint of clypeus (cly) and pronotum (pro) covered with soggy clay. (c–f) occlusion of the orifice shown in Figure 12a after end of rainfall, by injection of soggy clay (captured images with the automatic camera Brinno timelapseTLC200 Pro).
Correlation between intense rainfalls (////) and temporary and simultaneous opening of 35 turrets, during 3 episodes, between July 19 and 23, 2016.
When rainfalls are heavy, the soil becomes very wet and the moisture level in the burrow increases dramatically. In addition, rainwater in some places passes through a rapidly decomposing litter [3]. We can therefore imagine that the level of CO2 also increases. The temporary opening of the turret would therefore lower these rates, as well as the concomitant increase in pressure. The turret would thus appear as a regulating device of the physical–chemical conditions prevailing inside the burrow.
From the end of July begins the period of moults. The nymphs emerge from their turret after having opened it at the top, moult into a winged imago, on their own turret (Figure 14a,b) or on neighbouring vegetation (Figure 14c), then fly away for a brief adult life, during which males and females will mate.
(a) Exuvia attesting for the moulting of a nymph on its own building (photos Vanessa Gama). The white line represents the trip of the nymph between the opening at the top of the turret and the place where the moult took place. Note the rotation made by the nymph at the bottom of its turret. (b) imago from another nymph drying its wings after moulting on its own building. (c) Migration of a nymph from its turret to the branch where it has moulted. o: Opened turret.
We have identified three opening modes (Béguin, Gama and Ribamar Mesquita Ferreira, to be published), each accomplished using the hook-shaped end of the forelegs. It is therefore important to note that the opening behaviours of the top of the turret before moulting are totally distinct from those described above for the temporary opening of the top during intense rainfalls, and which are practiced by liquefaction of the top with soaked clay, without ever involving the front legs. For the opening of its turret, the nymph therefore has a register of behaviours specific to the function performed.
The most common method of opening before moulting (named program A) is worth explaining here. It takes place in two phases: a draft of the opening (Figure 15a–c), then an equalisation until the opening is remarkably circular (Figure 15d).
Four stages of opening a turret. (a) Beginning of the draft work (6:21 a.m.). (b) draft in progress (9:58 a.m.). (c) Draft completed (10:43 a.m.). (d) Equalisation of the border completed (11:35 a.m.). Observation on 19.08.2019.
Continuous monitoring makes it possible to describe in detail the behaviour of the nymph which creates the roughing by enlarging (Figure 16a–c) the initial perforation (Figure 15a) by a scraping carried out with the chitins’ end of its forelegs, up to obtain an elongated opening with irregular edges (Figure 16c). Working from the inside, the nymph is almost invisible. When the roughing is complete, the nymph appears (Figure 16c) and undertakes the work of equalisation, which is accomplished by scraping the edge with the chitins’ end of its forelegs (fl) also (Figure 16d), at the same time as it performs rotations. During this phase, the nymph’s head is clearly visible, and one can easily recognise its antennae (ant), front legs (fl) and clypeus (cly). Once the equalisation is complete (Figure 16e), the nymph disappears until its exit (Figure 16f) which takes place with a remarkable precision between 6 p.m. and 6:10 p.m. (local time). The 6 pictures in Figure 17 illustrate the rotations that the nymph performs at the same time as it levels the edge of the opening with its forelegs.
(a) 6:59 a.m.: Opening resulting from the enlargement of a small perforation, such as that shown on Figure 15a. (b) 7:03 a.m.: Continuation of the enlargement; the nymph is discernible. (c) 8:44 a.m.: The enlargement is finished; the nymph appears, on can distinguish its eyes (ey) and its clypeus (cly). (d) 9:38 a.m.: The nymph, gripped at the opening, is equalising the edge by scratching with the hook-shaped end of its forelegs (fl); one distinguishes also an antenna (ant) and the clypeus (cly). (e) 10:05 a.m.: The equalisation of the edge is complete, the opening is perfectly circular and the nymph withdrew. (f) 6:08 p.m.: The nymph leaves its building. Observation on 01.09.2019.
Six snapshots illustrating the nymph’s rotations when equalising the edge of the opening with its forelegs. (a) 9:15 a.m. (b) 9:35 a.m. (c) 9:37 a.m. (d) 9:38 a.m. (e) 9:41 a.m. (f) 9:45 a.m.. Arrows represent the rotations from the previous snapshot; ant: Antenna, fl: Foreleg, cly: Clypeus. Observation on 01.09.2019.
The monitoring of two individuals, from the exit of the nymph from its turret, until the flight of the imago, made possible to establish a detailed time schedule (Table 1).
Moulting period | Late July - early September (maximum frequency during the first week of August) |
---|---|
First perforation for the opening of the top of the turret | Before 7:00 a.m. |
End of opening (before equalising the edge) - program A | Between 8:30 and 11:00 a.m. |
Equalisation of the edge completed - program A | Between 10:00 and 12:00 a.m. |
Exit of the nymph through the opening at the top of its turret | Between 6:00 and 6:10 p.m. |
Immobilisation before moulting | Between 6:45 and 7:00 p.m. |
Downtime to moult | About 2 hours |
Downtime after moulting (drying the wings) | About 5 hours |
Taking flight | After 2 a.m. |
Work schedule of the nymph during its moult to imago.
Each time an edifice of G. chlorogena was discovered on the site we studied, an arborescent Fabaceae of the genus Tachigali has been identified in the vicinity (Béguin, Gama and Ribamar Mesquita Ferreira, to be published). The building of the nymph is usually the much farther from the tree the larger this one is. This fact is not surprising when one takes into account: 1°) that a cicada nymph can plant its tiny rostrum [4] to suck up the elaborate sap on which it feeds, only in radicular extremities of comparable size, close to the meristems; 2°) that the root ends of trees, in the Amazon rainforest where the layer of fertile soil is very small (of the order of 1 meter), instead of plunging in depth, spread out [5], forming a “root disc” whose radius is all the greater as the size of the tree is large.
During the 2 years 2018 and 2019, we identified 132 buildings of G. chlorogena distributed among 14 trees of the genus Tachigali; 11 from the species chrysophyllum and 3 from the species venusta. We therefore measured, for each building, the distance separating it from the tree2, as well as the diameter of the trunk of this latter3.
The results were reported on two graphs relating the diameter of a tree with: a) the distances to all the buildings associated with it (Figure 18a); b) the average distance to them (Figure 18b). The trend lines calculated with Excel by the smallest squares indicate clearly, on each of the graphs, a proportionality between the diameter of the trunk of the tree and its distance to a building of G. chlorogena, thereby providing a logical-mathematical support to the association between the nymphs of G. chlorogena and these Fabaceae4. G. chlorogena apparently being the main, if not the only, beneficiary of this relationship, we admit that this is a case of commensalism [6].
Relationship between Tachigali’s trunk diameters and distances to G. chlorogena’s edifices. (a) Plotting distances to all the buildings associated with a given tree. (b) Plotting the average distance to the buildings associated with a given tree.
The graph with plotting of all distances (Figure 18a) shows that part of the buildings are located near the trunk. We will be re-examining our statistics to determine if the location of the buildings is related to the Tachigali species (chrysophyllum or venusta). One thing is obvious; nymph buildings (i.e. nymphs feeding) close to the trunk involve root ramifications down to the level of the meristems from the proximal roots, particularly the buttresses. We recently discovered, at another place in the Adolfo Ducke reserve, a Tachigali station of a species (yet to be identified) which is neither chrysophyllum nor venusta, with nymph buildings (the species to which their hosts belong has not yet been confirmed as G. chlorogena) close to the trunk. In Santarém (Pará, Brazil), edifices of Fidicina chlorogena, a taxon since identified with G. chlorogena [1] have been described [7] near the base of the trunk; the species name of the tree was not reported. The same disposition was observed [8] for Orialella aerizulae (Boulard), which could also feed on Tachigali sp. as well as possibly on Tabebuia sp. This aspect of the dependency relationship between cicadas and their foster trees has to be explored in more detail.
The nymphs spend several years underground, moving to reach the fine roots (less than 2 mm) from which they suck, with their rostrum, the elaborate sap on which they feed (see above). They move forward by digging galleries with their forelegs, throwing the excavated material behind them.
On can consider [9] the beginning of the reproductive cycle at the time of mating (Figure 19a), after the nymph has emerged from its burrow and moulted into an imago (Figure 19h,i). The winged adults, stimulated by singing males, mate in trees. Then the females search for a stem in which they will plant their ovipositor several times [10] to deposit their eggs under the bark (Figure 19b). The larvae hatch on the twigs and then fall to the ground (Figure 19c) where they sink (Figure 19d). No one has yet observed their behaviour during this first phase of underground life. No one either knows the duration of the underground life until the exit to moult into a winged imago; it is estimated at several years.
Life cycle of G. chlorogena. (a) Mating. (b) Lay eggs. (c) After hatching, the tiny larvae (~ 2 mm) fall to the ground. (d) They bury themselves in the ground. (e) The nymph digs its gallery at the same time as it feeds on the sap of the roots. (f) As it moves forward and back, the nymph changes direction until it is vertically oriented. (g) The nymph digs the well which is a vertical gallery. (h) the turret is built from the well. (i) The nymph comes out of the turret to turn into an imago.
From excavations undertaken after moulding with cement (see above), which revealed an inclination of the deepest zone of the well, as well as in its vicinity traces of oblique galleries of intermediate inclinations (Figure 5b), one can envisage the upwards digging of a vertical well from a deep horizontal gallery (Figure 19e) by digging successive oblique galleries more and more inclined (Figure 19f).
The digging of deep horizontal galleries giving access to fine roots that the nymphs can use to feed themselves (see above), is consistent with the presence of a significant biomass of fine roots in the deep soil layer of the Amazon rainforest, as recent research has shown [11]. The digging of successive oblique galleries more and more inclined implies a capacity of the nymph to back up into an already dug burrow. Such a capacity is observed when one surprises, by the withdrawal of its turret, a nymph near the exit of its well. It can also be observed in experiments where the nymph is placed in a glass tube.
The key role mentioned above of urine in G. chlorogena, has been underlined in connection with the construction of the turret and the various maintenance activities described; in experiments where the turret was replaced by a glass tube [2], the observation was made of the rise of the urine soaked mass of clay that the nymph loads on its head. The role of urine, in European species of cicadas, was demonstrated [12] for digging underground burrows; abdominal gutters redirect urine to the anterior part of the body and allow the nymph to use it to soften the soil with its forelegs. In the case of G. chlorogena, urine is used to soften the clay [2]. Urine from cicadas also contains mucin, a glycoprotein, which strengthens the walls of burrows after drying [13].
Different reptiles, such as Lacertilians (Figure 20a), can be supposed as predators when a nymph opens or leaves its building. However ants, known by the vernacular name of Legion ants, or Formigas de coreção according to the local name, are the most dangerous (Figure 20b–e). They invade (Figure 20e) towers which have been opened (Figure 20b) following heavy rainfalls (see above: the role of the turret), and even manage to perforate the walls (Figure 20c). Entering a building, they devour the host, leaving only the chitins’ organs, such as the hook-shaped ends of the forelegs (Figure 20d).
(a) Lacertilian on an open turret. (b) Perforations (p) following passage of Legion ants on an open turret after an episode of heavy rainfall; o: Opening in the shape of its clypeus practiced by the nymph in response to rainfall. (c) Perforations (p) following a passage of army ants on a turret that the nymph closed with a clay occlusion (clc) after the cessation of precipitation. (d) Bottom of a well after passage of ants; the remainder of the hook of the end of a front leg (fl) is visible. (e) Ant in the bottom of a well.
This article has presented the activities of the nymph of the cicada G. chlorogena during its last year of larval life, in relation to its burrow, the visible part of which is a clay turret built from a well whose depth can reach one meter. The clay with which the turret is built probably comes from the base of the well, from the volcanic bedrock on which the fertile soil rests. The nymph mixes this clay with its urine and transfers the mixture to its forelegs and then to its clypeus, through its abdominal and thoracic “gutters” [9]. It can then climb this mortar to the surface and build its turret. By replacing a turret with a glass tube, it was possible [2] to observe the rise of a nymph with a mass of soggy clay on its clypeus.
The existence of a one meter deep well, dug vertically from the bottom thanks to a process of verticalization of a horizontal gallery (Figure 19f), validates the representation of the digging by the nymph of paths below the root base of the tree with which it is associated, in order to find appropriate roots to introduce its rostrum and feed on elaborated sap [10], which is confirmed, as mentioned above, by the recent demonstration [11] of a significant biomass of fine roots in the deep soil layer of the Amazon rainforest. To dig, the nymph shovels with its chitins’ forelegs, tears off earthy fragments which it impales on the bristles of its clypeus and deposits behind it [12]. It thus advances in a short gallery, which it opens in front of it and closes behind (Figure 19e). One question remains to be clarified; at what moment, and consecutively to which signals (external and/or endogenous), does the nymph begin its process of verticalization.
The well therefore has the status of a gallery, from which it is however distinguished by the fact that, after closing off the bottom, it is not closed as the nymph digs it, vertically and from bottom to top, until reaching the ground surface. The turret, for its part, is an additional device. Maintained sealed by the nymph (Figure 8), it appears to be devoted to maintaining appropriate conditions for the survival of the nymph in its burrow (see above; Role of the turret). As already mentioned, a minimum height of the turret is required by the nymph, which repairs or rebuilds it if necessary.
A behaviour, considered as motor coordination, involves [14, 15] stimuli (external and/or endogenous), as well as a recognition mechanism which is a neuronal structure. The observed richness of the nymph’s behaviours is therefore concomitant with an important perceptual component in its nerve system. The realisation of similar tasks (opening the top of the turret, for example) by different behaviours according to the circumstance, can be considered as falling under an elementary cognitive system. The execution, without learning, of complex motor sequences, such as the opening of the top of the turret before moulting, according to a straight defined time schedule, is the result of highly perfected innate programming.
The dependence of G. chlorogena on arboreal Fabaceae of the genus Tachigali is a very important aspect of its reproductive cycle. In this regard, an important question arises: do the very young larvae which have just burrowed (Figure 19b), have the same mode of nutrition? If so, are they also related to Tachigali to find fine root ends near the soil surface?
Many questions still remain open about cicadas, insects as popular as their biology is poorly understood.
I wish to express my gratitude to Prof. Ennio Candotti, who entrusted me with this research and constantly supported me, to Vanessa Gamma and José Ribamar Mesquita Ferreira for technical assistance, to Dr. Nállarett Dávila for the determination of the Tachigali, and to Olga Béguin for critical reading. The work was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNpq), grants no 385612/2014-1 et no 310012/2015-5, by the Fundação de Amparo à Pesquisa do Estado do Amazonas (FAPEAM), grant no 462/2018, and by Museu da Amazônia (MUSA), 69060-060 Manaus, AM, Brasil.
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\n\nSljedeća terminologija odnosi se na Odredbe i uvjete, te na sve naše ugovore:
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\n\nStranke, strane odnosi se na klijenta i na nas, ili samo na klijenta ili nas.
\n\nSve odredbe koje se odnose na ponudu, prihvat ili razmatranje plaćanja, a za koja mi pružamo asistenciju klijentu, bilo na ugovoreni ili fiksni način, a s ciljem da se ostvare potrebe i želje klijenta u svezi s našim uslugama, su podložne zakonskim odredbama Ujedinjenog Kraljevstva.
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