Blattabacterium Genome: Structure, Function, and Evolution Austin Alleman
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Feeding and Foraging Behaviors of Subterranean Termites (Isoptera:Rhinotermitidae)
Louisiana State University LSU Digital Commons LSU Historical Dissertations and Theses Graduate School 1989 Feeding and Foraging Behaviors of Subterranean Termites (Isoptera:Rhinotermitidae). Keith Scott elD aplane Louisiana State University and Agricultural & Mechanical College Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_disstheses Recommended Citation Delaplane, Keith Scott, "Feeding and Foraging Behaviors of Subterranean Termites (Isoptera:Rhinotermitidae)." (1989). LSU Historical Dissertations and Theses. 4838. https://digitalcommons.lsu.edu/gradschool_disstheses/4838 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Historical Dissertations and Theses by an authorized administrator of LSU Digital Commons. For more information, please contact [email protected]. INFORMATION TO USERS The most advanced technology has been used to photograph and reproduce this manuscript from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely afreet reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. -
Serial Horizontal Transfer of Vitamin-Biosynthetic Genes Enables the Establishment of New Nutritional Symbionts in Aphids’ Di-Symbiotic Systems
The ISME Journal (2020) 14:259–273 https://doi.org/10.1038/s41396-019-0533-6 ARTICLE Serial horizontal transfer of vitamin-biosynthetic genes enables the establishment of new nutritional symbionts in aphids’ di-symbiotic systems 1 1 1 2 2 Alejandro Manzano-Marıń ● Armelle Coeur d’acier ● Anne-Laure Clamens ● Céline Orvain ● Corinne Cruaud ● 2 1 Valérie Barbe ● Emmanuelle Jousselin Received: 25 February 2019 / Revised: 24 August 2019 / Accepted: 7 September 2019 / Published online: 17 October 2019 © The Author(s) 2019. This article is published with open access Abstract Many insects depend on obligate mutualistic bacteria to provide essential nutrients lacking from their diet. Most aphids, whose diet consists of phloem, rely on the bacterial endosymbiont Buchnera aphidicola to supply essential amino acids and B vitamins. However, in some aphid species, provision of these nutrients is partitioned between Buchnera and a younger bacterial partner, whose identity varies across aphid lineages. Little is known about the origin and the evolutionary stability of these di-symbiotic systems. It is also unclear whether the novel symbionts merely compensate for losses in Buchnera or 1234567890();,: 1234567890();,: carry new nutritional functions. Using whole-genome endosymbiont sequences of nine Cinara aphids that harbour an Erwinia-related symbiont to complement Buchnera, we show that the Erwinia association arose from a single event of symbiont lifestyle shift, from a free-living to an obligate intracellular one. This event resulted in drastic genome reduction, long-term genome stasis, and co-divergence with aphids. Fluorescence in situ hybridisation reveals that Erwinia inhabits its own bacteriocytes near Buchnera’s. Altogether these results depict a scenario for the establishment of Erwinia as an obligate symbiont that mirrors Buchnera’s. -
Isoptera Book Chapter
Isoptera 535 See Also the Following Articles Biodiversity ■ Biogeographical Patterns ■ Cave Insects ■ Introduced Insects Further Reading Carlquist , S. ( 1974 ) . “ Island Biology . ” Columbia University Press , New York and London . Gillespie , R. G. , and Roderick , G. K. ( 2002 ) . Arthropods on islands: Colonization, speciation, and conservation . Annu. Rev. Entomol. 47 , 595 – 632 . Gillespie , R. G. , and Clague , D. A. (eds.) (2009 ) . “ Encyclopedia of Islands. ” University of California Press , Berkeley, CA . Howarth , F. G. , and Mull , W. P. ( 1992 ) . “ Hawaiian Insects and Their Kin . ” University of Hawaii Press , Honolulu, HI . MacArthur , R. H. , and Wilson , E. O. ( 1967 ) . “ The Theory of Island Biogeography . ” Princeton University Press , Princeton, NJ . Wagner , W. L. , and Funk , V. (eds.) ( 1995 ) . “ Hawaiian Biogeography Evolution on a Hot Spot Archipelago. ” Smithsonian Institution Press , Washington, DC . Whittaker , R. J. , and Fern á ndez-Palacios , J. M. ( 2007 ) . “ Island Biogeography: Ecology, Evolution, and Conservation , ” 2nd ed. Oxford University Press , Oxford, U.K . I Isoptera (Termites) Vernard R. Lewis FIGURE 1 Castes for Isoptera. A lower termite group, University of California, Berkeley Reticulitermes, is represented. A large queen is depicted in the center. A king is to the left of the queen. A worker and soldier are he ordinal name Isoptera is of Greek origin and refers to below. (Adapted, with permission from Aventis Environmental the two pairs of straight and very similar wings that termites Science, from The Mallis Handbook of Pest Control, 1997.) Thave as reproductive adults. Termites are small and white to tan or sometimes black. They are sometimes called “ white ants ” and can be confused with true ants (Hymenoptera). -
Table S4. Phylogenetic Distribution of Bacterial and Archaea Genomes in Groups A, B, C, D, and X
Table S4. Phylogenetic distribution of bacterial and archaea genomes in groups A, B, C, D, and X. Group A a: Total number of genomes in the taxon b: Number of group A genomes in the taxon c: Percentage of group A genomes in the taxon a b c cellular organisms 5007 2974 59.4 |__ Bacteria 4769 2935 61.5 | |__ Proteobacteria 1854 1570 84.7 | | |__ Gammaproteobacteria 711 631 88.7 | | | |__ Enterobacterales 112 97 86.6 | | | | |__ Enterobacteriaceae 41 32 78.0 | | | | | |__ unclassified Enterobacteriaceae 13 7 53.8 | | | | |__ Erwiniaceae 30 28 93.3 | | | | | |__ Erwinia 10 10 100.0 | | | | | |__ Buchnera 8 8 100.0 | | | | | | |__ Buchnera aphidicola 8 8 100.0 | | | | | |__ Pantoea 8 8 100.0 | | | | |__ Yersiniaceae 14 14 100.0 | | | | | |__ Serratia 8 8 100.0 | | | | |__ Morganellaceae 13 10 76.9 | | | | |__ Pectobacteriaceae 8 8 100.0 | | | |__ Alteromonadales 94 94 100.0 | | | | |__ Alteromonadaceae 34 34 100.0 | | | | | |__ Marinobacter 12 12 100.0 | | | | |__ Shewanellaceae 17 17 100.0 | | | | | |__ Shewanella 17 17 100.0 | | | | |__ Pseudoalteromonadaceae 16 16 100.0 | | | | | |__ Pseudoalteromonas 15 15 100.0 | | | | |__ Idiomarinaceae 9 9 100.0 | | | | | |__ Idiomarina 9 9 100.0 | | | | |__ Colwelliaceae 6 6 100.0 | | | |__ Pseudomonadales 81 81 100.0 | | | | |__ Moraxellaceae 41 41 100.0 | | | | | |__ Acinetobacter 25 25 100.0 | | | | | |__ Psychrobacter 8 8 100.0 | | | | | |__ Moraxella 6 6 100.0 | | | | |__ Pseudomonadaceae 40 40 100.0 | | | | | |__ Pseudomonas 38 38 100.0 | | | |__ Oceanospirillales 73 72 98.6 | | | | |__ Oceanospirillaceae -
A Nondichotomous Key to Protist Species Identification of Reticulitermes
SYSTEMATICS A Nondichotomous Key to Protist Species Identification of Reticulitermes (Isoptera: Rhinotermitidae) 1 J. L. LEWIS AND B. T. FORSCHLER Department of Entomology, 413 Biological Sciences Building, University of Georgia, Athens, GA 30602 Ann. Entomol. Soc. Am. 99(6): 1028Ð1033 (2006) ABSTRACT A key was developed using morphological and behavioral characters to identify nine genera and 13 species of protists found in the hindgut of three Reticulitermes speciesÑ Reticulitermes flavipes (Kollar), Reticulitermes virginicus (Banks), and Reticulitermes hageni BanksÑby using the online IDnature guides by Discover Life. There are seven characters and 13 taxa, each attached to species descriptions, digital stills, or movies to aid in protist species identiÞcation. We chose characters for protist species identiÞcation that were easy to observe with live samples and a light microscope at 400ϫ magniÞcation. All 11 protists from R. flavipes and nine each in R. virginicus and R. hageni were recognized using original and revised species descriptions. This was the Þrst report of the protist genera Trichomitus from both R. virginicus and R. hageni. KEY WORDS symbiotic protists, termite identiÞcation, anaerobic protists identiÞcation, Parabasa- lia, Oxymonadida The anaerobic symbiotic protist orders found in the (workers have Ϸ57 Ϯ 11%), because it is not found in hindgut of lower termites (Isoptera) include Tricho- R. virginicus and R. hageni (Lewis and Forschler monadida Kirby, Oxymonadida Grasse´, and Hyper- 2004b). Trichonympha agilis Leidy (1877) is Ϸ19 Ϯ 5% mastigida Grassi & Foa` (Yamin 1979). None of these of the protist population in R. virginicus compared protist species are found outside of the insect host with 4 Ϯ 3% in R. -
Parallel and Gradual Genome Erosion in the Blattabacterium Endosymbionts of Mastotermes Darwiniensis and Cryptocercus Wood Roaches
GBE Parallel and Gradual Genome Erosion in the Blattabacterium Endosymbionts of Mastotermes darwiniensis and Cryptocercus Wood Roaches Yukihiro Kinjo1,2,3,4, Thomas Bourguignon3,5,KweiJunTong6, Hirokazu Kuwahara2,SangJinLim7, Kwang Bae Yoon7, Shuji Shigenobu8, Yung Chul Park7, Christine A. Nalepa9, Yuichi Hongoh1,2, Moriya Ohkuma1,NathanLo6,*, and Gaku Tokuda4,* 1Japan Collection of Microorganisms, RIKEN BioResource Research Center, Tsukuba, Japan 2Department of Life Science and Technology, Tokyo Institute of Technology, Tokyo, Japan 3Okinawa Institute of Science and Technology, Graduate University, Okinawa, Japan 4Tropical Biosphere Research Center, University of the Ryukyus, Okinawa, Japan 5Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Prague, Czech Republic 6School of Life and Environmental Sciences, University of Sydney, NSW, Australia 7Division of Forest Science, Kangwon National University, Chuncheon, Republic of Korea 8National Institute for Basic Biology, NIBB Core Research Facilities, Okazaki, Japan 9Department of Entomology, North Carolina State University, Raleigh, North Carolina, USA *Corresponding authors: E-mails: [email protected]; [email protected]. Accepted: May 29, 2018 Data deposition: This project has been deposited in the International Nucleotide Sequence Database (GenBank/ENA/DDBJ) under the accession numbers given in Table 1. Abstract Almost all examined cockroaches harbor an obligate intracellular endosymbiont, Blattabacterium cuenoti.Onthebasisof genome content, Blattabacterium has been inferred to recycle nitrogen wastes and provide amino acids and cofactors for its hosts. Most Blattabacterium strains sequenced to date harbor a genome of 630 kbp, with the exception of the termite Mastotermes darwiniensis (590 kbp) and Cryptocercus punctulatus (614 kbp), a representative of the sister group of termites. Such genome reduction may have led to the ultimate loss of Blattabacterium in all termites other than Mastotermes. -
Ancestry and Adaptive Radiation of Bacteroidetes As Assessed by Comparative Genomics
1 Ancestry and adaptive radiation of Bacteroidetes as assessed by comparative genomics 2 3 Raul Munoza,b,*, Hanno Teelinga, Rudolf Amanna and Ramon Rosselló-Mórab,* 4 5 a Department of Molecular Ecology, Max Planck Institute for Marine Microbiology, D-28359 6 Bremen, Germany. 7 b Marine Microbiology Group, Department of Ecology and Marine Resources, Institut Mediterrani 8 d’Estudis Avançats (CSIC-UIB), E-07190 Esporles, Balearic Islands, Spain. 9 10 * Corresponding authors: 11 Raul Munoz, Marine Microbiology Group, Carrer Miquel Marquès 21, 07190 Esporles, Illes 12 Balears, Spain. e-mail: [email protected] 13 Ramon Rosselló-Móra, Marine Microbiology Group, Carrer Miquel Marquès 21, 07190 Esporles, 14 Illes Balears, Spain. e-mail: [email protected] 15 16 17 18 Keywords + 19 Bacteroidetes, Na -NQR, alternative complex III, caa3 cytochrome oxidase, gliding, T9SS. 20 21 Abbreviations 22 m.s.i.: median sequence identity. 23 24 25 26 27 ABSTRACT 28 As of this writing, the phylum Bacteroidetes comprises more than 1,500 described species with 29 diverse ecological roles. However, there is little understanding of archetypal Bacteroidetes traits on 30 a genomic level. We compiled a representative set of 89 Bacteroidetes genomes and used pairwise 31 reciprocal best match gene comparisons and gene syntenies to identify common traits that allow to 32 trace Bacteroidetes’ evolution and adaptive radiation. Highly conserved among all studied 33 Bacteroidetes was the type IX secretion system (T9SS). Class-level comparisons furthermore 34 suggested that the ACIII-caa3COX super-complex evolved in the ancestral aerobic bacteroidetal 35 lineage, and was secondarily lost in extant anaerobic Bacteroidetes. -
Table S5. the Information of the Bacteria Annotated in the Soil Community at Species Level
Table S5. The information of the bacteria annotated in the soil community at species level No. Phylum Class Order Family Genus Species The number of contigs Abundance(%) 1 Firmicutes Bacilli Bacillales Bacillaceae Bacillus Bacillus cereus 1749 5.145782459 2 Bacteroidetes Cytophagia Cytophagales Hymenobacteraceae Hymenobacter Hymenobacter sedentarius 1538 4.52499338 3 Gemmatimonadetes Gemmatimonadetes Gemmatimonadales Gemmatimonadaceae Gemmatirosa Gemmatirosa kalamazoonesis 1020 3.000970902 4 Proteobacteria Alphaproteobacteria Sphingomonadales Sphingomonadaceae Sphingomonas Sphingomonas indica 797 2.344876284 5 Firmicutes Bacilli Lactobacillales Streptococcaceae Lactococcus Lactococcus piscium 542 1.594633558 6 Actinobacteria Thermoleophilia Solirubrobacterales Conexibacteraceae Conexibacter Conexibacter woesei 471 1.385742446 7 Proteobacteria Alphaproteobacteria Sphingomonadales Sphingomonadaceae Sphingomonas Sphingomonas taxi 430 1.265115184 8 Proteobacteria Alphaproteobacteria Sphingomonadales Sphingomonadaceae Sphingomonas Sphingomonas wittichii 388 1.141545794 9 Proteobacteria Alphaproteobacteria Sphingomonadales Sphingomonadaceae Sphingomonas Sphingomonas sp. FARSPH 298 0.876754244 10 Proteobacteria Alphaproteobacteria Sphingomonadales Sphingomonadaceae Sphingomonas Sorangium cellulosum 260 0.764953367 11 Proteobacteria Deltaproteobacteria Myxococcales Polyangiaceae Sorangium Sphingomonas sp. Cra20 260 0.764953367 12 Proteobacteria Alphaproteobacteria Sphingomonadales Sphingomonadaceae Sphingomonas Sphingomonas panacis 252 0.741416341 -
The Phylogeny of Termites
Molecular Phylogenetics and Evolution 48 (2008) 615–627 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev The phylogeny of termites (Dictyoptera: Isoptera) based on mitochondrial and nuclear markers: Implications for the evolution of the worker and pseudergate castes, and foraging behaviors Frédéric Legendre a,*, Michael F. Whiting b, Christian Bordereau c, Eliana M. Cancello d, Theodore A. Evans e, Philippe Grandcolas a a Muséum national d’Histoire naturelle, Département Systématique et Évolution, UMR 5202, CNRS, CP 50 (Entomologie), 45 rue Buffon, 75005 Paris, France b Department of Integrative Biology, 693 Widtsoe Building, Brigham Young University, Provo, UT 84602, USA c UMR 5548, Développement—Communication chimique, Université de Bourgogne, 6, Bd Gabriel 21000 Dijon, France d Muzeu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, 04263-000 São Paulo, SP, Brazil e CSIRO Entomology, Ecosystem Management: Functional Biodiversity, Canberra, Australia article info abstract Article history: A phylogenetic hypothesis of termite relationships was inferred from DNA sequence data. Seven gene Received 31 October 2007 fragments (12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase I, cytochrome oxidase II Revised 25 March 2008 and cytochrome b) were sequenced for 40 termite exemplars, representing all termite families and 14 Accepted 9 April 2008 outgroups. Termites were found to be monophyletic with Mastotermes darwiniensis (Mastotermitidae) Available online 27 May 2008 as sister group to the remainder of the termites. In this remainder, the family Kalotermitidae was sister group to other families. The families Kalotermitidae, Hodotermitidae and Termitidae were retrieved as Keywords: monophyletic whereas the Termopsidae and Rhinotermitidae appeared paraphyletic. -
Universidad De Guadalajara
2005 B - 2010 A COD. 005216222 Universidad de Guadalajara CENTRO UNIVERSITARIO DE CIENCIAS BIOLÓGICAS Y AGROPECUARIAS -,......-. ( ... - .-'' (. ~· .. - -. ....__, ' .,- .. ~ , , ~ .. ... ~ GUIA DIDACTICA PARA EL ESTUDIO DE TERMITAS ~.. ~ TRABAJO DE TITULACIÓN EN LA MODALIDAD DE PRODUCCIÓN DE MATERIALES EDUCATIVOS QUE PARA OBTENER EL GRADO DE LICENCIADO EN BIOLOGÍA PRESENTA GIUSEPPE RAFAEL GALINDO RODRÍGUEZ DIRECTOR: DR. FLORENCIO RECENDIZ HURTADO Las Agujas Zapopan, Jalisco. Junio de 201 1. 1 Universidad de Guadalajara Centro Universitario de Ciencias Biológicas y Agropecuarias Coordinación de Carrera de la Licenciatura en Biología COORD-810-090/2011 C. GIUSEPPE RAFAEL GALINDO RODRÍGUEZ PRESENTE Manifestamos a usted, que con esta fecha, ha sido aprobado su tema de titulación en la modalidad de PRODUCCION DE MATERIALES EDUCATIVOS opción PAQUETE DIDÁCTICO con el titulo: "Guía didáctica -para el estudio de termitas'', para obtener la Licenciatura en Biologia. Al mismo tiempo le infon11amos, que ha sido aceptado como director de dicho tra'bajo al Dr. Florencia Recendiz Hurtado. Sin más por el momento, aprovecha111os para enviarle un cordial saludo. ATENTAMENTE "PIENSA Y TRABAJA" Las Agu1as, Nextipac, Za popan. Jal , 01 de junio de 2011. DR . ESA E ZES ESQUIVIAS PRESIDENTE DEL COMITÉ DE TITULACIÓN # M.C. GLORIA PARADA BARRERA SECRETARIO DEL COMITÉ DE TITULACIÓN Las Agujas, Zapopan, Jalisco, México. C.P. 45110. AP 39-82. Tels. fíll-.111177711 ~íl 1hR?017d .... 1 1?<:.1 i:-~v ~7T7l 1 ~o FORMAF Dra. Teresa de Jesús Aceves Esquivias. Presidente del Comité de Titulación. Licenciatura en Biología. CUCBA. Presente Nos permitimos informar a usted que habiendo revisado el trabajo de titulación, modalidad Producción de materiales educativos, opción Paquete didáctico con el título: "Guía didáctica para el estudio de termitas" que realizó el pasante Giuseppe Rafael Galindo Rodríguez con número de código 005216222, consideramos que ha quedado debidamente concluido, por lo que ponemos a su consideración el trabajo terminado para autorizar su impresión y presentación. -
Morphometric Analysis of Coptotermes Spp. Soldier Caste (Blattodea: Rhinotermitidae) in Indonesia and Evidence of Coptotermes Gestroi Extreme Head-Capsule Shapes
insects Article Morphometric Analysis of Coptotermes spp. Soldier Caste (Blattodea: Rhinotermitidae) in Indonesia and Evidence of Coptotermes gestroi Extreme Head-Capsule Shapes Bramantyo Wikantyoso 1,2,*, Shu-Ping Tseng 3, Setiawan Khoirul Himmi 2 , Sulaeman Yusuf 2 and Tsuyoshi Yoshimura 1 1 Research Institute for Sustainable Humanosphere (RISH), Kyoto University, Gokasho, Uji, Kyoto 611-0011, Japan; [email protected] 2 Research Center for Biomaterials, Indonesian Institute of Sciences (LIPI) Jl. Raya Bogor km 46 Cibinong, Bogor 16911, Indonesia; [email protected] (S.K.H.); [email protected] (S.Y.) 3 Department of Entomology, University of California, 900 University Avenue, Riverside, CA 92521, USA; [email protected] * Correspondence: [email protected] Simple Summary: The morphological characteristics of the soldier caste in termites provide valuable taxonomic information at the species level. Head-shape variation in soldiers was often used as an indicative characteristic in some genera. While species with egg-shaped and waterdrop-shaped head capsule (HC), Coptotermes gestroi and C. curvignathus, respectively, are familiar in Indonesia, neither a measurement nor head index may avoid the subjectivity of shape interpretation. We conducted linear Citation: Wikantyoso, B.; Tseng, S.-P.; and geometric morphometrics analyses of soldiers’ HC of Coptotermes spp. obtained from various Himmi, S.K.; Yusuf, S.; Yoshimura, T. locations in Indonesia. Although subtle differences were observed, the posterior parts of the HC Morphometric Analysis of laterally expanded in a gradual manner in C. gestroi, C. sepangensis, and C. curvignathus in that order. Coptotermes spp. Soldier Caste Furthermore, three extreme head-shape variations of C. -
The Cockroach Blattella Germanica Obtains Nitrogen from Uric Acid
Physiology The cockroach Blattella germanica obtains nitrogen from uric acid through a rsbl.royalsocietypublishing.org metabolic pathway shared with its bacterial endosymbiont Research Rafael Patin˜o-Navarrete1, Maria-Dolors Piulachs2, Xavier Belles2, Cite this article: Patin˜o-Navarrete R, Piulachs Andre´s Moya1, Amparo Latorre1 and Juli Pereto´1,3 M-D, Belles X, Moya A, Latorre A, Pereto´ J. 1 2014 The cockroach Blattella germanica obtains InstitutCavanilles de Biodiversitat i Biologia Evolutiva, Universitat de Vale`ncia, C/Catedra`tic Jose´ Beltra´nn8 2, Paterna 46980, Spain nitrogen from uric acid through a metabolic 2Institut de Biologia Evolutiva (Consejo Superior de Investigaciones Cientı´ficas and Universitat Pompeu Fabra), pathway shared with its bacterial Passeig Marı´tim de la Barceloneta n8 37-49, Barcelona 08003, Spain endosymbiont. Biol. Lett. 10: 20140407. 3Departament de Bioquı´mica i Biologia Molecular, Universitat de Vale`ncia, Burjassot 46100, Spain http://dx.doi.org/10.1098/rsbl.2014.0407 Uric acid stored in the fat bodyof cockroaches is a nitrogen reservoir mobilized in times of scarcity. The discovery of urease in Blattabacterium cuenoti,theprimary endosymbiont of cockroaches, suggests that the endosymbiont may participate Received: 21 May 2014 in cockroach nitrogen economy. However, bacterial urease may only be one piece in the entire nitrogen recycling process from insect uric acid. Thus, Accepted: 2 July 2014 in addition to the uricolytic pathway to urea, there must be glutamine synthe- tase assimilating the released ammonia by the urease reaction to enable the stored nitrogen to be metabolically usable. None of the Blattabacterium genomes sequenced to date possess genes encoding for those enzymes.