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An Introduction to the Microscopical Study of Diatoms Robert B. McLaughlin Edited by John Gustav Delly & Steve Gill An Introduction to the Microscopical Study of Diatoms An Introduction to the Microscopical Study of Diatoms Robert B. McLaughlin Edited by John Gustav Delly and Steve Gill Editorial note: Many readers will recognize the name of Robert B. ―Mac‖ McLaughlin as the author of a regular Diatom column published in The Microscope during the 1980‘s and 1990‘s. He also wrote two books in the Microscope Series on Accessories and Contrast Methods. He was a contributor to the diatom literature, including numerous papers printed in the Quekett Microscopical Club Journal and, indeed, has had a diatom named for him, Gomphonema mclaughlinii (Reichardt). Over 50 years ago, while still in Alaska, and prior to his work on the diatoms of the Kenai Deposit, Mac started to write a diatom textbook for himself acting more as a compiler or organizer of material from, generally, older, scarcer diatom works. He was, in his words, like a fish out of water, and wished to self-educate so as to be able to do meaningful work with the microscope and diatoms. He read and studied extensively, initially for his own amusement and edification, and succeeded to a remarkable degree. Then, one day, thinking that he might help others by letting them know what he had learned were the important things to study, where to find information and publications, and, in short, distill his ―Collecting and hard-won knowledge, he started ―the book‖. For preparing diatoms, as well as disclosures illustrations he wanted to use figures from Friedrich about examination and Hustedt‘s publication, Vom Sammeln und präparien culture methods.‖ der Kieselalgen sowie angaben uber Untersuchungs und Kulturmethoden-aberholden, (ABB.XI T.4, S.I., 1929); the publisher Cramer had told him that the work was long out-of-print, and that there would be no problem about using illustrations from it. Mac‘s book eventually came to be over 525 typed papers, while still needing some appendices, index, table of contents, revision, and editing. More than 25 years ago, a publisher told Mac that they wanted to prepare his book for distribution, but, incredulously, they made no progress over a ten year period! Mac was, understandably, disheartened by the whole venture, and the book was put away; in addition, he had moved to Santa Fe. The matter of the book was brought up at The Hooke College of Applied Sciences, and it was decided that it would be a valuable contribution to make this knowledge available by some means for those wishing to start the microscopical study of diatoms. As it turned out, Mac was by then an octogenarian, his eyesight was poor, and reading a chore. He had donated his equipment, samples, and references to the California References to Hustedt Academy of Sciences to be with the donations from in the main body of the Hanna. Being thus unable to do anything more with book should assume the book, he gave permission to the American Editor that the detail referred at Hooke College of Applied Sciences to do with it as to may be in any of his he saw fit. After much discussion, it was decided to volumes. Page i Robert B. McLaughlin abandon search for an editor or anybody else to attempt to complete it, and to publish it online, as it is, even to include Mac‘s own sketches and handwritten notes. There is precedent for this tack, as in the re-publication of Mann‘s The Bullet’s Flight, with Pope‘s hand-written marginalia, and Lee‘s Note-Book of an Amateur Geologist. Mac no longer had the Hustedt book he intended to use for illustrations, but readers will find adequate illustrative material in Hustedt‘s three-part Die Kieselalgen or in Schmidt‘s three-volume Atlas der Diatomaceen-Kunde, both of which have been reprinted. Thus, the Hooke College of Applied Sciences feels that rather than let Mac‘s 525+ page manuscript on diatoms, descend, unseen, into oblivion, it would be better to transcribe it as best we can and let it stand on its own merits. Whilst it is true that some of the text and conjecture is now dated and superseded, the majority of the content is as useful to both amateur and professional as it was when Mac first wrote it. The manuscript was in a semi-edited state. Where Mac authorized any correction it has been included, simple typographical errors have been corrected, any other editing has been at the discretion of the Editors, including the addition of an extensive bibliography. The layout was designed for US Royal sized paper with a view to it being printed. The formatting of the document reflects this intention. Space has been left in the margins for user‘s own notes. American Editor: John Gustav Delly (Scientific Advisor to Hooke College of Applied Sciences, Westmont, Illinois, U.S.A.) British Editor: Steve Gill (Co-Editor of The Amateur Diatomist, United Kingdom) Winter 2012 Many of the chemical procedures described are included in a historical context. Though some might still be used by trained and suitably equipped chemists it is not recommended that others use the procedures or techniques described without first fully understanding the dangers and ramifications of misuse etc. Should you choose to embark on any of the procedures then you do so at your own risk and the Editors will not and cannot be held responsible. Page ii An Introduction to the Microscopical Study of Diatoms Remembering Robert B. “Mac” McLaughlin 1922 – 2012 During the time of the final editorial process it was learned that Mac had died (on 6th April 2012) having turned 90 years old in January of that year. A brief biographical note follows: Mac, as he was known to his friends, will be remembered by long-time readers of The Microscope as the author and editor of the popular "Diatoms" column, which appeared in each quarterly issue for 10 years in 1985-1995. He also wrote two highly respected books in the Microscope Series: Accessories for the Light Microscope (1975) and Special Methods in Light Microscopy (1977). In addition, he authored or co-authored numerous articles on diatoms for specialty journals, including a major contribution (with John L. Stone), ―Some Late Pleistocene Diatoms of the Kenai Peninsula, Alaska‖ (Nova Hedwigia, 1986). Mac‘s contribution to the diatom and microscopy literature was formally recognized when he received the Annual Award of the State Microscopical Society of Illinois. The ultimate recognition of Mac as a diatomist came when he had a diatom named after him: Gomphonema mclaughlinii E. Reichardt. He proudly displayed a photomicrograph of this diatom on the letterhead of his stationery. Incidentally he was also mentioned by name in a crime novel by Patricia Cornwell as an expert in diatom identification during a forensic examination of trace evidence. Mac had received his Federal Communications Commission assigned Radio Operator License in 1938, when he was 16 years old. He built his own radio station, of course, and was assigned the call sign W6QIN. In 1940, Mac joined the United States Navy for six years, and at boot camp in San Diego, California, he qualified for Aviation Radio School. He was subsequently sent to the Naval Air Station at Seattle, Washinton, where he became a radio operator aboard a PBY aircraft. The squadron went to Kodiak, Alaska, in early summer 1941. It was here that he became a high-speed Navy telegrapher. The Navy did not furnish bugs (a kind of telegraph key with springs and adjustable weights that allows for Page iii Robert B. McLaughlin adjustment of speed and touch) at the time, and Mac supplied his own - a MacKey. Mac‘s ―fist‖ was beautiful; his tone was pure, and his well-formed characters were characteristically recognizable - it was a pleasure to copy his transmissions. After Pearl Harbor, Mac and a buddy of his volunteered for a mission to install a one receiver/one transmitter radio station for weather reporting at Chernofski Bay on the other end of Unalaska Island. After a year at Umnak, Mac was transferred to be in charge of the Amchitka Transmitter Station at Kirilof Point. Then it was back to the States, where he became Chief-in-Charge of the Naval Transmitter Station at Pensacola, Florida. In 1946, Mac went to Tri-State College in Angola, Indiana, and graduated with a Bachelor of Science degree in Electrical Engineering. He then moved to Anchorage, Alaska and the Civil Aeronautics Administration, where he was responsible for both the Intra-Alaska and Trans-Oceanic sites. In 1961, Mac became interested in diatoms, and started his studies on the diatoms found in Alaska‘s Kenai Peninsula. It was during this time that he started his unpublished 500-page personal manuscript notebook on diatom structure and identification, which is the subject of this book. His letters, which were always typed on his ―mill‖ (manual typewriter), and his Morse code transmissions were always full of fascinating detail about his diatom studies. A selection of these communications was shown to the Editor of The Microscope at the time, and Mac was invited to write a regular column on diatoms. Fortunately, he accepted, and provided 10 years of articles and two books. Mac‘s microscopy lab in Santa Fe, which was housed in his radio shack, was equipped with a Leitz CM microscope and a Zeiss Standard microscope, Interestingly, while in Santa Fe, Mac became a Research Associate of the Museum of New Mexico, where, from 1981 to 1991, he specialized in the microscopical examination of folk and fine art. One project he worked on involved the question of the origin of a specific piece of Southwest Indian pottery. Mac sampled the clay used to make the pottery, and found diatoms mixed with the clay. He identified the diatom flora, and then proceeded to examine clays from possible sites. He ended up pinpointing the exact origin of the clay, and thus, the tribe responsible for making that particular pottery piece. Successes like these gave him great satisfaction. As a side interest, Mac enjoyed collecting full plate blocks of postage stamps, and, of course, stamps that depicted microscopes. When Mac started to have trouble with his eyes, he replaced his Zeiss photomicrographic camera with a video unit, so that he could view his images on a TV monitor. When it became apparent that his failing eye-sight would no longer permit him to do serious diatom work, he donated his reference collection of more than 4,000 diatom reference slides, books, and other equipment to the California Academy of Sciences, so that they could be added to what G Dallas Hanna had already established at the Academy. Mac, however, retained one microscope and about 200 diatom slides and spent his final days recording notes on these 200 slides so that his daughter Susan might have them; she has every intention of using her father‘s microscope and viewing the specimens while reading his notes. He will be missed by all those who enjoyed his diatom publications and also by his many friends across the world. Page iv An Introduction to the Microscopical Study of Diatoms Although “Mac” never saw this version of his manuscript we feel that had he done so and been able to see it to completion he would have added the following: For my daughter Susan and to beginning diatomists everywhere. Page v Robert B. McLaughlin Page vi An Introduction to the Microscopical Study of Diatoms CONTENTS Page Editorial Note i Remembering Robert B. ―Mac‖ McLaughlin iii List of Figures xvi List of Tables xix PART I 1 CHAPTER 1. 1 1. MORPHOLOGY . 1 1.1. Structure of the Cell Wall 1 1.2. General Structure 1 1.3. Symmetry 3 1.3.1. Line-Symmetrical Frustules 6 1.3.2. Plane-Symmetrical Frustules 11 1.4. Girdle Bands 13 1.5. Intercalary Bands 14 1.6. Septa 18 1.7. Craticular Plates 20 1.8. Liostephania 21 1.9. Valve Structure 22 1.9.1. Size 22 1.9.2. Shape 22 1.9.3. Striae, Punctae, and Pores 27 1.9.4. Areolae 29 1.9.5. Canaliculi 30 1.9.6. Costae 31 1.9.7. Cell Walls 32 1.9.7.1. Single-layer Cell Wall (Laminar type) 32 1.9.7.2. Two-layer Cell Wall (Loculate type) 33 1.9.8. Processes 35 1.9.9. Hyaline Areas 39 1.9.10. The Raphe 40 1.9.11. Special Morphological Nomenclature 47 1.10. Polymorphism 54 1.11. Abnormal Forms 55 1.12. Descriptive Terms and Features and Examples 55 1.13. The Cell Contents 57 1.13.1. The Elementary Plant Cell 57 1.13.2. The Diatom Cell 59 CHAPTER 2. 67 2. PHYSIOLOGY 67 2.1. Nutrition 67 2.2. Growth 68 2.3. Movement 72 2.4. Formation of the Cell Wall 75 Page vii Robert B. McLaughlin CHAPTER 3. 77 3. REPRODUCTION 77 3.1. Vegetative Cell Division 77 3.2. Auxospores 79 3.2.1. Normal Type A. 81 3.2.1.1. The gametes are undifferentiated or isogamous 81 3.2.1.2. Mother cell produces wandering and resting gamete 81 3.2.1.3. One mother cell produces two wandering gametes 81 3.2.1.4. The gametes behave according to no rule 81 3.2.2. Normal type B. 81 3.2.2.1. Four spermia are produced 81 3.2.3. Reduced Type A. 81 3.2.3.1. The gametes behave isogamously 82 3.2.3.2. The gametes behave anisogamously 82 3.2.4. Reduced Type B. 82 3.2.4.1. Two gametes of one mother cell copulate 82 3.2.4.2. The sexual nuclei of a mother cell copulate 82 3.2.5. Reduced Type C. 82 3.2.5.1. From one mother cell there develops two 82 3.2.5.2. From one mother cell there develops one 82 3.2.5.2.1. Parthenogenetically 82 3.2.5.2.2. Purely vegatatively 82 3.3. Resting Spores 83 3.4. Microspores 84 CHAPTER 4. 87 4. DISTRIBUTION AND ECOLOGY 87 4.1. Fresh and Brackish water Diatoms 87 4.1.1. Water Habitats 88 4.1.1.1. Lakes 88 4.1.1.2. Other Quiet Water Habitats 88 4.1.1.3. Rivers and Streams 89 4.1.1.4. Other Moving Water Habitats 89 4.1.1.5. Aerial Habitats 90 4.1.1.6. Chemical and Physical Factors 91 4.1.1.7. Geographic Distribution 93 4.2. Marine Diatoms 94 CHAPTER 5. 97 5. CLASSIFICATION 97 5.1. Mode of Growth 98 5.2. Size 99 5.3. Form of the Frustule 99 5.3.1. Zone 99 5.3.2. Symmetry or Asymmetry 99 5.3.3. Central Nodule 99 5.3.4. Raphe 100 5.3.5. Terminal Fissures 100 5.3.6. Axial and Central Areas 100 5.3.7. Longitudinal Lines 100 Page viii