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Invertebrate Embryology and Reproduction

Invertebrate Embryology and Reproduction

Professor in Department of Zoology, Faculty of Science, Alexandria University, Egypt

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To my dearly beloved father…

MAHMOUD EL-SAYED EL-BAWAB

Dedication

and my highly valued teacher and professor.

PROFESSOR DR. MAHMOUD MOHAMED RAMADAN

Classification

Body

Class

Class

Class Cubozoa

Class Anthozoa

Representatives

References

VI Phylum Ctenophores Definition

Triploblastica Acoelomata

VII Phylum

Turbellaria

VIII Phylum Nemertea

Pseudocoelomata

Phylum Mollusca

Preface

Invertebrate Embryology and Reproduction is an essentially descriptive book. Although this branch has long been a core subject recognized as a necessary and valuable part of Zoology and Biology courses, it was not considered as a definite course in Egypt until the 1970s.

To give a brief account of my academic background, I ought to start by saying that I was an extremely curious student, fascinated by embryology, with an ambition to pursue an academic research career. I graduated from the Faculty of Science, Alexandria University in June 1955; at that time, unfortunately there were no available posts, and I was not able to join the faculty to fulfil my dream. However, I joined the Oceanography department to pursue my postgraduate studies. The Dean of the Faculty of Science at the time, the late Professor Doctor Abdel Fattah Mohammad, who happened to be teaching a course for this programme, became familiar with my work, and was quick to recognize my devotion to academe. Consequently, he offered me a post in the Zoology department; thus I became a university staff member in January 1956, and continue to be one. As a demonstrator in the practical laboratories, I was exposed to a variety of specialties in the science of zoology. However, I was always interested in embryology, and intended to pursue research in experimental embryology, but due to the lack of financial resources, my request to equip a suitable laboratory was rejected.

I was selected by the head of the Zoology department then, the late Prof. Dr Mahmoud Ramadan, who was the Professor of Invertebrates, as the demonstrator for the invertebrate laboratories. Consequently, my research was primarily concerned with invertebrates. My MSc and PhD theses were on Crustacea, and the PhD was on spermatophore, under the supervision of Prof. Ramadan. I feel privileged to have been his postgraduate student, and to have had the opportunity to work closely with him. His indelible influence on me, and on many others, is eternal. On the other hand, Prof. Dr Samy Riad, as a joint supervisor of my PhD, was the one who planned the course of the work, and basically taught me the principles of research, and the importance of integrity in carrying it out. I was incessantly guided and motivated by his discussions and decisive criticism. I was also well supported by the capable senior colleagues at this stage, Prof. Anga Ramadan and Prof. Milad Ishaa.

Eventually, I was considered the successor of Prof. Ramadan in Invertebrates. To follow on from his work, I created the ‘Invertebrate school’ in the Zoology department. Later, as a professor, I could confidently describe myself as an untraditional academic, working hand in hand with my students. I persistently examined the specimens myself, and aided in the actual writing of the theses, and in the interpretations and discussions.

By the time I had finished my PhD and started working as a lecturer on Invertebrates in the Zoology department at the Faculty of Science in Alexandria University, Oceanography had just started to be included as an undergraduate course. I was recommended by the head of the Zoology department at the time, the late Prof. Dr Shukralla Nakhla Sidra, to the head of the Oceanography department, Prof. Dr Saad El Wakil, to teach Invertebrate Embryology for the first time. I was the first, and the only one, to prepare and teach this course starting in 1970. I taught it to the general degree and special degree students, as well as for the pre-master’s students. I also taught Experimental Embryology to the master’s students. The great challenge then was that there were virtually no resources. I had to manage by using information out of the development and reproduction chapters from the available invertebrate books. For years, I had to ask any colleague or student travelling abroad to look for books specializing in invertebrate embryology.

It was only due to the genuine willingness of my departmental colleagues to help that I was supplied with the main available references in the field of invertebrate embryology over the years. The late Prof. Dr Mahmoud El-Zayat, Professor of Zoology, while in the United States ordered a book on invertebrate embryology edited by Kumé and Dan, translated from Japanese by J.C. Dan. After an unusually long wait, he had to place another order. Fortunately, I received both copies. Likewise, I can only be grateful to all other colleagues who travelled abroad over the years and went out of their way to provide me with the most-needed books: Dr Essam Abdulla, lecturer of Zoology, brought me Experimental Embryology of Marine and Fresh Water Invertebrates, by G. Reverberi, 1971, from the United States. Dr Desouky Mekawy, Professor of Zoology, brought me the valuable German book by O. Pflugfeder, 1970: Lehrbuch dez Entwicklungsgeschichte und Entwicklungsphysiologie der Teire. Dr Safaa Zaki, Lecturer of Zoology, brought

me an unpublished MSc thesis, from the University of Maryland, United States, titled A Histological Study of the Development of the Ovary and Accessory Reproductive Organs of the Blue Crab, Callinectes sapidus (Rabhdum), by L.E. Cronim, 1942. Dr Mohga El Abbady managed to send me different articles from the Congress Library in the United States, including an article about Helix Pomatia (E. Lind, 1973). Locally, people were equally supportive. Dr Mahmoud Ibrahim, Assistant Prof. of Molecular Biology, was always keen to provide me with all the related articles he found while searching on the Internet, in addition to his continual assistance with any article related to molecular biology.

The library in the Faculty of Science in Alexandria University has always offered me a fantastic service in terms of facilitating book borrowing. The late Prof. Dr Ibrahim El-Kholy, who was the Vice Dean of the Faculty of Science and Chair of the library, never rejected a purchase request that I submitted as a lecturer in Zoology. I therefore managed to get to the library a few purchases, including the series of Reproduction of Marine Invertebrates (volumes I–V). However, this series was not complete. I searched for it myself in UK bookshops during a visit there and realized that this series was not in circulation any more. I wrote to one of the editors, Giese, asking for the rest of the series, and received a reply from Pearse, the co-editor, who sent me volume VI, Echinodermata and Lophophorates, as well as all the related articles.

All those difficulties encouraged me into the idea of writing a comprehensive book about invertebrates. Having gone deeper in embryology through teaching, I found that books on the subject are quite rare in the market. For years, the update of the invertebrate embryology was confined to limited information from references dealing with general invertebrate reproduction. It did not cover all the phyla, and

the books never included animals from the Mediterranean region. This does not fulfil the need for a comprehensive and exclusive update of the field. Even the recent publications, Embryology, Constructing the Organism, by Gilbert and Raunio, 1997, and Developmental Biology by Gilbert, 2000, are general and neither is exclusive to invertebrate embryology. In addition, the Atlas of Comparative Invertebrate Embryology by Dr Martin E.L. Scriba contains figures and not text. Its aim is to show clear figures for the invertebrate embryology, but not to describe and discuss the developmental stages for further research and teaching purposes. I therefore decided that it would be a pleasure and honour to write a comprehensive book as an Egyptian scholar, and to invest all the great effort spent preparing that course over the years.

This book therefore gathers all the information dealing with descriptive invertebrate embryology and updates this information in a single source, as a basic reference for teaching courses and research. The difficulty of obtaining teachable or research data in the field of descriptive embryology, not only in the Egyptian libraries but everywhere, demonstrated the need for a new work that collates all the available data concerning that field. The collection herein is sought from specialized references, in addition to general textbooks in reproduction, carried out by a professional expert in that field. It includes the description of embryology of newly studied species in most of the phyla, together with their systematic positions, and the results obtained from different researches using advanced techniques and different procedures, especially electron microscopical ones. It is specific to descriptive invertebrate embryology, including numerous invertebrate species. I hope that this effort will provide a coherent integrated account of the field that I attempted to cover, to the satisfaction of the reader.

Acknowledgements

It is only with the support of my family that this book was accomplished: both emotional support and unstinting encouragement as well as actual involvement with the work. I am immensely proud of my grandchildren, Ziyad and Zeinah Kalati, for their contribution in formatting the figures and revisions. The same goes to my daughter, Dr Siham Abdel Fattah, for her vital involvement with the work. My appreciation goes to my youngest daughter, Dr Azza Abdel Fattah, for the time that she generously invested in this work. She was involved in all stages of the work, and her thorough and meticulous revisions were invaluable. She handled all the required correspondence with the publisher, as well as the overwhelming copyright permissions requirements. It is in fact thanks to her zeal and keenness that this work was completed. I would like to thank all who helped me during the planning and writing of this book, even the small efforts, which

collectively made this project possible. I am truly grateful to Dr Amal Khafage, Lecturer of Invertebrates in the National Institute of Oceanography and Fisheries, who took time from her busy schedule to help in the searches for related articles to the points suggested. My sincere appreciation goes to Mr Mohammad Abdou, my nephew, who helped with the figure plates and Photoshop work, and to Dr Tarek Adnan, whose work in Bibliotheca Alexandria facilitated my obtaining copies of difficult-to-get articles. Finally, I would like to thank the staff at Elsevier for their patience and their help and expertise in editing and production.

Last, but not least, my sincere thanks to all members of the Department of Zoology, Faculty of Science, Alexandria University, for being a genuine family to me, sharing love, support, and care throughout my long journey of working in the department.

Chapter 1

Historical review

An invertebrate (nonvertebrate) refers specifically to an animal without a backbone, in contrast to the vertebrates represented by fish, amphibians, reptiles, birds, and mammals. An invertebrate animal has no central skeleton and has a ventral nervous system, while the heart is dorsal in position, with an open circulatory system, and its integument is formed of secretory substances, generally tanned proteins and sometimes chitinized.

At present, about 1.5 million species of invertebrates have been named and about 8000–10,000 new species are added to the list each year. With the exception of the protozoans (about 30,000 species) and the vertebrates (about 43,000 species), all other animals are invertebrates, comprising altogether about 95% of the animals in the animal kingdom. Strictly speaking, the protozoans are also invertebrates, but a distinction is sometimes made. If we also consider the extinct species, the total number of species that have lived would be much greater than that of the existing forms alone. The numbers of species are just estimations and vary enormously amongst zoologists. Evolution of new species is a continuous process; in addition, names of new species added annually to the list might result from the discovery of forms that have long been in existence but have only just come to light.

The invertebrates are grouped into about 30 phyla, the number of which varies among different zoologists. A phylum is somewhat difficult to define, but it may be considered as a basic structural pattern of evolutionary descent from a common ancestry, with its members more or less homogeneous and bearing a unique combination of distinct characteristics. The members of a phylum are all constructed along the same general plan. Some of the phyla are major ones (according to the constituting number of species), while others are minor phyla: a minor phylum can be a group separated (e.g. by class and/or order) from a major phylum such as Nemertinia from Platyhelminthes.

Embryology and Reproduction. https://doi.org/10.1016/B978-0-12-814114-4.00001-1

Nemertinia was elevated to the rank of a phylum since its members have additional and/or different characters. A minor phylum can also be a newly discovered group of animals, which was established independently as a phylum, e.g. phylum Pogonophora.

According to embryology, the invertebrate phyla are grouped into two lines:

Annelid line Echinoderm line

Egg potency Mosaic Equipotent

Cleavage Spiral Radial

Blastomeres Determinate Indeterminate Gastrulation Delamination Invagination

Coelom Schizocoele Enterocoele

Site of mouth Protostome Deuterostome

Larva Trochophore (ciliated band around the equator) Pluteus (ciliated band around the mouth)

Embryology (from Greek έμβρύον, embryon, ‘unborn embryo’; and -λογίa, -logia). It may be defined as that branch of biological science that deals with the development of the individual organisms. It is concerned with successive series of changes, in form and function, through which the initial germ of the new individual is transformed into a sexually mature adult. Normally, the germ with which development commences is an egg that has been fertilized by a sperm. The sexually mature adult is an individual that has developed to a point where it can produce mature eggs if a female, or sperm if a male. Sometimes the word ontogeny is used as a synonym for embryology, but more often it is defined to include the entire life history of an individual from its origin to its death.

Many principles of embryology apply to both invertebrate animals and vertebrates. Therefore, the study

of invertebrate embryology has advanced that of vertebrate embryology, despite the presence of many differences. The most obvious is the release of a larva in numerous invertebrate species before the completion of development.

The history of embryology has passed through three phases: fact-finding or descriptive, comparative, and experimental embryology. More recently, the molecular studies have been established.

DESCRIPTIVE PHASE

The first famous name associated with this period is Aristotle.

Aristotle (384–322 BC)

With the possible exception of Plato, Aristotle is by far the most influential philosopher in the history of Western thinking. Aristotle studied natural sciences, and these studies dominated until early modern times. Aristotle’s influence was vast and extended far beyond philosophy. For example, he was the founder of biology, and Charles Darwin (1809–82) regarded him as the most important contributor to more than one subject.

The influence of Aristotle’s family (his father was a physician) initiated his strong interest in anatomy, and the structure of living organisms in general. He described the reproduction and development of many kinds of animals. He was the first to study and write the accounts of development of the hen’s egg. His Macedonian origin, being a student, and then a teacher in Plato’s academy, and his connections with the group of outstanding scientists surrounding Plato (who worked in a wide variety of subjects, ranging from medicine and biology to mathematics and astronomy), contributed to him developing remarkable observation skills. These media had greatly influenced and characterized the scientific thinking of Aristotle. He was able to reach and formulate some scientifically significant findings as follows:

1. He noted the different methods by which animals originate from eggs, and described them as oviparous, viviparous, and ovoviviparous animals, and divided them into the following categories:

● Mammals: because their young arose from a mixture of male and female fluids and are born alive. The eggs of mammals had not been discovered at that time.

● Sharks: arose from eggs that are retained within the body of the mother and are also born alive.

● Reptiles and birds: in which the eggs are complete ones.

● Amphibia and bony fishes: in which the eggs are incomplete (complete eggs are those with an albumen coat and a shell).

2. He identified the two major cell division patterns by which embryos are formed: complete division of the egg

(holoblastic), as in frogs and mammals; and incomplete division of the egg (meroblastic), as in chicks.

3. He was the first to figure out the functions of the placenta and the umbilical cord.

4. He also observed that the cephalic (anterior) part of the embryo grows faster than the posterior one (Fig. 1.1C and D). Therefore he discovered—as pointed out by Joseph Needham (one of the modern embryological pioneers) in 1934—the morphogenetic cephalocaudal gradient.

5. As a result of describing different embryos, he claimed the most valuable theory, which initiated massive arguments and discussions during later times, the epigenesis. This theory states that development proceeds from a simple formless beginning to the complex organization of the adult.

After Aristotle, there was hardly any progress in embryology for the next 2000 years. It was not until the 15th and 16th centuries that the science in general, and embryology in particular, was refreshed. The most famous man during that period was William Harvey.

The English Physician William Harvey (1578–1657) (Before the invention of the microscope)

Harvey described the development of the chick in great detail. He also contributed a study of the development of the deer, and compared the two with each other. He was the first to observe the blastoderm of the chick embryo—that small region of the egg that contains the yolk-free cytoplasm which gives rise to the embryo. He was also the first to notice that islands of blood cells form before the heart does (Fig. 1.1A and B). Harvey suggested that the amniotic fluid might function as a shock absorber for the embryo. His most important finding was his conclusion that all animals arise from eggs, but this was based on theoretical considerations. He famously stated, ‘Omne vivum ex ovo’. He finally published a book, following his numerous studies and results, Exercitationes de generatione animalium (On Animal Generation), published in 1651. His doctrine of omne vivum ex ovo (all life comes from the egg) was the first definite statement against the idea of spontaneous generation.

In addition to all this work, Harvey’s main line of research was dealing with the blood circulation in the human body, and he became an embryologist, in addition to being the founder of modern physiology and medicine. He published his theories in a book titled Exercitatio Anatomica de Motu Cordis et Sanguinis in Animalibus (An Anatomical Study of the Motion of the Heart and of the Blood in Animals), where he explained how the heart propelled the blood in a circular course through the body. His discovery was received with great interest in England, although it was greeted with some scepticism on the Continent (www. en.wikipedia.org/wiki/William_Harvey and www.bbc.co.uk/history/historic_figures/harvey_william. shtml).

Auditory vesicle

Aor tic arches

Atrium Hear t

Tr uncus

ar terious

Ventricle

Liver

r udiment

Forelimb bud

Somite

Hindlimb bud

Myelencephalon

Metencephalon

Mesencephalon

Diencephalon

Choroid fissure

Lens Eye

Sensor y retina

Pigmented retina

Olfactory pit

Telencephalon

Vitelline vein

Vitelline ar tery

Tail

FIG. 1.1 Depictions of chick developmental anatomy. (A) Dorsal view (looking ‘down’ at what will become the back) of a 2-day chick embryo, as depicted by Marcello Malpighi in 1672. (B) Ventral view (looking ‘up’ at the prospective belly) of a chick embryo at a similar stage, seen through a dissecting microscope and rendered by F.R. Lillie in 1908. (C) Eduard D’Alton’s depiction of a later stage 2-day chick embryo in Pander (1817). (D) Modern rendering of a 3-day chick embryo after Carlson (1981). Details of the anatomy will be discussed in later chapters. (From Gilbert (2000).)

Discovery of the microscope

Leeuwenhoek, Anton van [lay'-vuhn-hook, ahn' -tohn vahn] (1632–1723)

Leeuwenhoek was a Dutch biologist and microscopist. He was a businessman, and became interested in science when he developed an interest in lenses and started building

simple microscopes as a hobby. Each microscope consisted of a flat brass or copper plate in which a small, single glass lens was mounted. The lens was held up to the eye, and the object to be studied was placed on the head of a movable pin, just on the other side of the lens. Leeuwenhoek built over 400 microscopes, whose focuses were amazingly accurate and clear. Many of these microscopes still exist. The most

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Sailings, 3

Seasons, the, 20

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Time diagram, 51

Tropical year, 66 V

Vega as a pole star, 31

Vernal equinox, 20 dates of, 67

Vertical circles, 12

Weir’s azimuth diagram, 98

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