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The Histology of the Third Instar Larva of the Apple Maggot, Rhagoletis Pomonella (Walsh)

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The Histology of the Third Instar Larva of the Apple Maggot, Rhagoletis Pomonella (Walsh) University of Massachusetts Amherst ScholarWorks@UMass Amherst Masters Theses 1911 - February 2014 1972 The histology of the third instar larva of the apple maggot, Rhagoletis pomonella (Walsh). John Knell University of Massachusetts Amherst Follow this and additional works at: https://scholarworks.umass.edu/theses Knell, John, "The histology of the third instar larva of the apple maggot, Rhagoletis pomonella (Walsh)." (1972). Masters Theses 1911 - February 2014. 3015. Retrieved from https://scholarworks.umass.edu/theses/3015 This thesis is brought to you for free and open access by ScholarWorks@UMass Amherst. It has been accepted for inclusion in Masters Theses 1911 - February 2014 by an authorized administrator of ScholarWorks@UMass Amherst. For more information, please contact [email protected]. jf-om THE HISTOLOGY OF THE THIRD INSTAR LARVA OF THE APPLE MAGGOT, RHAGOLETIS POMONELLA (WALSH) A Thesis Presented By John Knell Submitted to the Graduate School of the University of Massachusetts in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Sept. 1972 Major Subject: Entomology THE HISTOLOGY OF THE THIRD INSTAR LARVA OF THE APPLE MAGGOT, RHAGQLETIS POMONELLA (WALSH) A Thesis By John Knell Approved as to style and content by: J'hJUL (Member) Sept. 1972 ACKNOWLEDGEMENTS I wish to express my gratitude to my advisor Dr. J. G. Stoffolano whose patience and guidance were of great value in the completion of this thesis. Thanks also go to Drs. P. Barbosa, H. Potswald and * S. Wasti who furnished my with many valuable histological techniques and Dr. K. Trammel of the New York Agricultural Experiment Station at Geneva, New York for supplying me with apple maggot pupae. Funds for this study were provided through Hatch grant #3^2 awarded to Dr. J. G. Stoffolano. iii TABLE OF CONTENTS Page I. INTRODUCTION. 1 II. LITERATURE REVIEW A. Integument. 3 B. Anal organ... 5 C. Tracheal system. 6 D. Alimentary tract. 7 E. Fat body.... 9 F. Nervous system 1. Central nervous system. 10 2. Stomatogastrie system. 13 3. Sense organs... 14 G. Imaginal discs... 1? H. Reproductive system... 19 I. Circulatory system.*. 20 J. Endocrine system... 22 K. Excretory system... 23 L. Bacterial symbiotes.- 25 III. MATERIALS AND METHODS A. Apple maggot colony.. 26 B. Histological techniques 1. Killing, relaxation and fixation... 28 2. Dehydration, clearing, infiltrating and embedding... 28 iv 3. Sectioning and staining. 30 4. Gross dissections.. 30 5. Microscopy and photomicroscopy. 31 IV. RESULTS AND DISCUSSION A. Integument. 32 B. Anal organ... 36 C. Tracheal system..... 38 D. Alimentary tract. 42 E. Fat body. 53 F. Nervous, system 1. Central nervous system... 55 2. Stomatogastric system,...... 62 3. Sense organs. 64 G. Imaginal discs. 70 H. Reproductive system...... ?6 I. Circulatory system...... 80 J. Endocrine system. 86 K. Excretory system. 89 L. Symbiotes. 94 V. SUMMARY. 96 VI. CONCLUSIONS. 97 VII. LITERATURE CITED. 98 VIII. PLATES. 108 v I. INTRODUCTION Since its original description under the name Trypeta pomonella by Walsh (186?), the apple maggot, Rhagoletis pomonella has remained an orchard pest of some importance in much of the U. S. and southeastern Canada, In temperate climates the apple maggot is univoltine. Adults emerge from late June through early August (Rivard, 1968) and apparently feed on aphid honeydew adhering to the surfaces of apple leaves and fruit (Neilson and Wood, I966). After a pre-oviposition period of about 12 days, the females mate and begin to lay eggs singly, just beneath the skin of the fruit with the aid of a sharp ovipositor. The larvae tunnel through the fruit spreading the bacterium Pseudomonas melophthora which causes a brown rot to develop in the apple flesh. Growth of both larvae and bacteria is slow until the fruit ripens and drops after which the apple maggot develops rapidly, leaves the rapidly decomposing apple and pupates in the soil. Winter is passed in the pupal stage, a number of pupae diapausing for two years. Due to the detrimental environmental effects of chemical insecti¬ cides, it would seem that future emphasis must be placed on alternative control measures. Many of these will depend on exploitation of certain aspects of the biology of target species, thus making basic research in this area desirable. The higher flies (cyclorrhapha) include a number of important pest species and often (for instance, the apple maggot) the larva is the important stage. However, little basic work has been done with maggots other than Drosophila, Musca and large Calliphorids, espe¬ cially with regard to histology. In fact, so far only the work of Perez 1 2 (1910) on Calliphora has dealt with the normal histology of the organ systems of the entire animal. The present study will attempt to rectify this by presenting a description of the histology of a typical cyclor- rhaphan larva, the apple maggot. Since maggots are very similar morphologically, this work will miti¬ gate interpretation of histological sections of the larvae of most cyclorrhapha. A knowledge of the appearance of the normal tissues would thus allow recognition of pathological conditions brought about through a biological control such as a pathogen or parasite. The major damage to fruit inflicted with apple maggot is actually caused by its associated symbiote, P. melophthora. This bacterium is in¬ troduced through oviposition and produces the brown rot always associated with apple maggot tunneling which eventually destroys the fruit. There¬ fore, a second objective of this study is to locate P, melophthora in the body of the animal and to determine if the larva contains any special structures for carrying its symbiotes into the pupal stage. In order to avoid the difficulty involved in working with the very small first and second instars, this study will concern only mature (third instar) larvae. II. LITERATURE REVIEW A. Integument The general organization of larval cyclorrhaphan cuticle into endocuticle and epicuticle has "been known since the work of Lowne (1890-92) on Calliphora. Dennell (1946) performed a variety of histo- chemical tests on the developing larval cuticle of Sarcophaga falculata and found that each of the two original layers could be further subdi¬ vided into two layers, the inner and outer endocuticle and the inner and outer epicuticle, Locke (1957) studied the epicuticle with the electron microscope and determined that the outer layer, which he termed the cuticulin, was present over most of the body surface of insects being absent only in some sense organs (Slifer, 1961) and the gut (Bertram and Bird, I96I). The cuticulin is extremely important as it essentially determines the surface characteristics of the cuticle as well as protecting the developing deeper layers from the potent molting fluid enzymes (Locke, 1966). The chemical composition of maggot cuticles was investigated by Pryor (1940) who was mainly interested in the localization of phenols during cuticle tanning. His histochemical tests on Calliphora showed phenol restricted to the epicuticle in the feeding larva. Fraenkel and Rudall (1940, 1947) studied the composition of the larval cuticles of Calliphora ervthrocephala and Sarcophaga falculata using a variety of chemical tests as well as X-ray diffraction and concluded that the larval cuticle is a polysaccharide-protein complex, 60% of which is chitin, and that in the larva these chitin filaments are oriented ran- 3 4 domly allowing flexibility. Richards (1956) performed microincinera¬ tion and elemental analysis studies on dried cuticles of Sarcophaga bullata and found traces of 25 elements left in the ash with magnesium being the predominant cation. The cuticulin layer left a relatively large amount of ash in which most of the iron content of the cuticle was * present. A number of authors (Dennell, 1946; Wolfe, 1954; Kennaugh, 1965* Filshie, 1970) have described pore canals from the larvae of cyclorrhaphous diptera, Locke (I96I) reported that no pore canals are present in the larva of Calpodes ethlius (Lepidoptera:Hesperiidae). 5 B. Anal organ This plate of thin endocuticle has been used by taxonomists for some time as a diagnostic character (Zimin, 1950 "but its function is still in doubt, El Shatoury (1955) claimed that in larval Drosophila the thin cuticle contracts rhythmically thus acting as an accessory % pulsating organ. Gloor (1949) found that the anal organ of Drosophila darkened after immersion in a dilute solution of silver nitrate indicating that this area is more permeable to ions than the surround¬ ing integument. On the basis of Gloor's work, Quintart (1961) proposed an osmoregulatory function for the anal organ, a view also held by Stoffolano (1970). 6 C. Tracheal system The most thorough study of the morphology of the tracheal system in larval Diptefa is that of Whitten (1955. 1957). Whitten's detailed (1955) paper deals comparatively with this system and reveals a common pattern occurring through the order which may still be recognized in the highly modified cyclorrhapha. Her later (1957) paper follows the development of the tracheal system of Drosophila through larva, pupa and adult. Recently, Whitten (1972) reviewed the morphology of the insect tracheal system. Keilin (1944) also studied the comparative morphology of this system in larval Diptera but was more concerned with the struc¬ ture, location, number and development of the spiracles. His lengthy paper includes a section on the paraspiracular (peristigmatic) glands in dipterous larvae. These glands were first reported by Leydig (1859). Gazagnaire (1886) noted the long, coiled duct in the gland of Eristalis tenax and rightly theorized its function to be to conduct a hydrophobic oil to the cuticle surrounding the spiracular openings. Keilin et al. (1935) found these glands in mosquito larvae and were able to discern oil droplets in the ducts leading to the spiracles. Bates (1934) described these glands in the anterior and posterior spiracles of the apple maggot while Phillips (1939) found them in the anterior spiracles of the walnut husk fly, Rhagoletis suavis.
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