Evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system

Summary A review is made of the ruminant digestive system in its morphophysiological variations and adaptations relating to foraging behaviour, digestive physiology, to interactions between plants and ruminants and to geographic and climatic diversity of ruminants' ecological niches. Evidence is provided for evolutionary trends from an extreme selectivity mainly for plant cell contents and dependence upon a fractionated fore- and hindgut fermentation, to an unselective intake of bulk roughage subjected to an efficient plant cell wall fermentation, mainly in the forestomachs. The review is based on detailed comparative morphological studies of all portions of the digestive system of 65 ruminant species from four continents. Their results are related to physiological evidence and to the classification of all extant ruminants into a flexible system of three overlapping morphophysiological feeding types: concentrate selectors (40%), grass and roughage eaters (25%) and intermediate, opportunistic, mixed feeders (35%). Several examples are discussed how ruminants of different feeding types are gaining ecological advantage and it is concluded that ruminants have achieved high levels of digestive efficiency at each evolutionary stage, (including well-documented seasonal adaptations of the digestive system) and that ruminant evolution is still going on. Deductions made from the few domesticated ruminant species may have, in the past, biased scientific evaluation of the free-ranging species' ecology. The main threat to a continuous ruminant evolution and diversity appears to be man's neglect for essential ecological interactions between wild ruminants and their specific habitats, which he alters or destroys.Abbreviations bw body weight - CS concentrate selector - DFC distal fermentation chamber (distended caecocolon) - GR grass and roughage eater - IM intermediate (mixed) feeder - PFC proximal fermentation chamber (ruminoreticulum/forestomachs) - RR Ruminoreticulum - SCFA Short-chain fatty acis (acetic, butyric, propionic acid set free by rumen bacteriae) - SE Surface enlargement (of absorptive mucosa) Supported by German Research Community grant DFG Ho 273/6Dedicated to Professor Dr. Dr. h.c. Dietrich Starck on the occasion of his 80th birthday     

Polygalacturonase activity and variation in ripening of papaya fruit with tissue depth and heat treatment

Effects of tissue position (viz. outer vs inner mesocarp) and heat treatment (48°C, 20 min) on variations in polygalacturonase (EC 3.2.1.15 and EC 3.2.1.67) activity and ripening of fruits of Carica papaya L. cv. Backcross Solo were investigated. Polygalacturonase activity increased during ripening concomitantly with an increase in tissue softness and soluble polyuronide level. Throughout ripening, inner mesocarp tissue was softer and contained higher polygalacturonase activity than outer mesocarp tissue. Titratable acidity as well as ß-galactosidase (EC 3.2.1.23) activity also increased during ripening; however, unlike polygalacturonase, their level or activity was lower in inner than in outer mesocarp. Ascorbic acid could partially account for the increase in titratable acidity during ripening but contributed very little to the differences in titratable acid levels between outer and inner mesocarp. Heat treatment had no effect on either fruit softness or titratable acidity, but it markedly reduced the increase in ascorbic acid and polygalacturonase activity during ripening. Ripening, as reflected by changes in tissue softness and polygalacturonase activity, progressed outwardly from the interior towards the exterior of the fruit. The effect of heat treatment in suppressing polygalacturonase activity was relatively greater in inner than in outer mesocarp, suggesting that sensitivity of the enzyme to heat treatment may vary with stage of ripeness of the tissue.     

Identification of cDNA clones for tomato (Lycopersicon esculentum Mill.) mRNAs that accumulate during fruit ripening and leaf senescence in response to ethylene

Changes in gene expression during foliar senescence and fruit ripening in tomato (Lycopersicon esculentum Mill.) were examined using in-vitro translation of isolated RNA and hybridization against cDNA clones.During the period of chlorophyll loss in leaves, changes occurred in mRNA in-vitro translation products, with some being reduced in prevalence, whilst others increased. Some of the translation products which changed in abundance had similar molecular weights to those known to increase during tomato fruit ripening. By testing RNA from senescing leaves against a tomato fruit ripening-related cDNA library, seven cDNA clones were identified for mRNAs whose prevalence increased during both ripening and leaf senescence. Using dot hybridization, the pattern of expression of the mRNAs corresponding to the seven clones was examined. Maximal expression of the majority of the mRNAs coincided with the time of greatest ethylene production, in both leaves and fruit. Treatment of mature green leaves or unripe fruit with the ethylene antagonist silver thiosulphate prevented the onset of senescence or ripening, and the expression of five of the seven ripening- and senescence-related genes.The results indicate that senescence and ripening in tomato involve the expression of related genes, and that ethylene may be an important factor in controlling their expression.Abbreviations cDNA copy-DNA - MW molecular weight - PAGE polyacrylamide gel electrophoresis - SDS sodium dodecyl sulphate     

Small mitochondrial DNA molecules of wild abortive cytoplasm in rice are not necessarily associated with CMS

Summary Mitochondrial DNA was isolated from leaf tissue of both the cytoplasmic male sterile line of Indica rice variety V41, which carries wild abortive (WA) cytoplasm, and from the corresponding maintainer line. In addition to the main mitochondrial DNA, four small plasmid-like DNA molecules were detected in both the male sterile and fertile lines. Restriction analysis of total mitochondrial DNA from the male sterile and fertile lines showed DNA fragments unique to each. Our findings suggest that the four small mitochondrial DNA (mtDNA) molecules are conserved when WA cytoplasm is transferred into different nuclear backgrounds. However, there is no simple correlation between the presence/ absence of small mitochondrial DNA molecules and the expression of WA cytoplasmic male sterility (CMS).     

Fruit photosynthesis

Abstract. In addition to photosynthesis as in the leaf, fruit possess a system which refixes CO₂ from the mitochondrial respiration of predominantly imported carbon. This pathway produces malate by the action of phosphoenolpyruvate carboxylase, PEPC, (E.C. 4.1.1.31) and appears to be regulated primarily by the cytosolic concentration of HCO₃/CO₂ and malate. Malate is stored in the vacuole as malic acid, constituting a major carbon pool and a potential substrate for respiration. The PEPC in apple fruit proves to be an efficient form of the enzyme with low Michaelis constants, i.e. Km = 0.09 mol m^-3 PEP and 0.2 mol m^–3 HCO₃, and large Ki= 110 mol m^-3 HCO³. In fleshy fruit, chlorophyll and chloroplasts are unevenly distributed; they resemble the C₃ sun-type and arc concentrated in the perivascular tissue, with smaller chloroplasts, fewer grana per chloroplast and a larger degree of vacuolation than commonly found in a leaf of the same species. Fruit photosynthesis often compensates for respiratory CO₂ loss in the light. However, due to respiration in the dark, CO₂ loss is in excess of photosynthetic gain in the light, such that a continual loss of CO₂ was observed in the diurnal cycle and which is maintained throughout fruit development. The rate of CO₂ exchange decreases on a fresh weight or surface basis, but increases with fruit ontogeny on a per fruit basis, causing accumulation of several percent CO₂ in the internal cavity. Stomata are present in the outer epidermis of those fruits examined, but with a 10-to 100-fold lesser frequency than in the abaxial epidermis of leaf of the same species. The number of Stomata is set at anthesis and remained constant, while the stomatal frequency decreases as the fruit surface expands. Stomata are as sensitive as in leaves in the early stages of fruit development, but often are transformed into lenticels during fruit ontogeny, thereby decreasing the permeability of the outer epidermis. The discrepancy between the CO₂-concentrating mechanism provided by PEPC analogous to C₄/CAM Photosynthesis and the kinetics of fruit PEPC, characteristic of C₃/non-autotrophic tissue, suggests the definition of a new type of ‘fruit photosynthesis’ rather than its categorization within an existing type.     

Electroantennogram responses of the Mediterranean fruit fly, Ceratitis capitata to identified volatile constituents from calling males

Fifty-six compounds from the odor of calling, sexually mature, laboratory reared males of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) were isolated by headspace trapping on Tenax columns and identified using GC/MS techniques (69 total compounds were detected). Electroantennogram responses (EAGs) to 54 of the 56 identified compounds as well as 5 analogs were tested on both sexes. Significant differences between the sexes in their responsiveness were found in 9 of the 54 identified compounds tested. There was no correlation between the amplitude of the EAG response and the relative abundance of compound identified from headspace analysis. Of the five major identified components, three elicited relatively small EAG responses, while two elicited large EAGs compared to the hexan-1-ol standard. The relative ranking of EAG responses were: methyl and ethyl hexenoates and hexanoates > C₄–C₆ esters and/or acetates > ethyl and methyl octenoates > monoterpenes > sesquiterpenes > C₂–C₅ acetates, alcohols and ketones. Behavioral bioassays on each of the five major identified components as well as a blend of six of the compounds showed some degree of attractancy to virgin females which in some cases approached the response to a pheromonal standard (male odors absorbed onto filter paper). These results are discussed in relationship to the insect's antennal sensitivity to putative pheromone components and/or allomonal components and to other reported C. capitata pheromone studies.

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Résumé Cinquante-six composés de l'odeur de mâles de C. capitata Weidemann, élevés en laboratoire, sexuellement mûrs et en appel, ont été isolés par piégeage sur colonnes tenax et identifiés par la technique GC/MS (69 composés avaient été détectés en tout). Les électroantennogrammes (EAGs) ont été examinés chez les deux sexes pour 54 des 56 composés identifiés et 5 de leurs analogues. Des différences significatives entre les sexes ont été observées pour 9 des 54 composés identifiés. Il n'y avait pas de corrélation entre l'ampleur de l'EAG et l'abondance relative du composé lors de son isolement. Pour les 5 principaux composés identifiés, 3 ont induit des EAGs relativement faibles, tandis que 2 étaient importants, par comparaison avec l'Hexane-1-ol utilisé comme témoin. Le classement relatif des EAG a été: hexénoates et hexanoates d'éthyl et de méthyl C₄–C₆ esters et/ou acétates octénoates d'éthyl ou de méthyl monoterpènes sesquiterpènes C₂–C₅ acétates, alcools et kétones. Les expériences de comportement avec chacun des 5 composés principaux identifiés, comme avec des mélanges de 6 composés ont mis en évidence une attraction des femelles vierges qui dans quelques cas avoisine la réponse à la phéromone témoin (odeur du mâle absorbée sur papier filtre). Ces résultats sont discutés en fonction de la sensibilité de l'antenne d'insexte aux composés supposés de la phéromone et aux composés allomonaux, et en fonction des autres études connues sur les phéromones de C. capitata.

     

A Markov approach for describing post-fire succession of vegetation

Vegetation dynamics in the coastal area of the Seto Inland Sea region in Japan, where wild fires occur frequently, were described using a stationary Markov model. In this region, vegetation types ofMiscanthus-Pleioblastus grassland,Lespedeza-Mallotus scrub,Pinus-Rhododendron forest andCrassocephalum-Erechtites community have been identified, and these show cyclic succession under the influence of fires. The model uses parameters determining fire frequency and rate of successional change to analyze the effect of variation in these parameters on the areal ratio of each vegetation type at equilibrium and on the time taken for one vegetation type to succeed another (elapsed successional time). The effect of fire frequency differs between hypothetical habitats with high and low productivity. A policy for vegetation management in areas of high and low productivity is proposed. The advantages and limitations of applying Markov models to studies of vegetation succession are also discussed.     

Auxins,ABA and gibberellin-like activity in abscising and non-abscising flowers and pods of Vicia faba L.

Abscission probability varies among floral positions within inflorescences of Vicia faba L. Flowers from proximal positions have a greater chance to develop into mature pods than flowers from more distal positions which normally abscise either as older flowers or as young pods. In three field experiments with the indeterminate single stem variety Herz-Freya, changes in the contents of extractable auxins, abscisic acid (ABA) and gibberellins in flowers and pods during their development, and their possible influence on abscission were investigated.Inflorescences at different positions along the stem were divided into the two proximal and the remaining fruits. The content of all three hormones was at a low level during flower development, increased greatly in parallel with dry matter accumulation in the young pods, and then decreased to maturity. The first hormone to increase in the fruits was auxin and this took place when abscission from the distal positions began. ABA and gibberellins at this time were still at a low level. This ontogenic course of hormone production was very similar in fruits of both positions within an inflorescence, but in flowers and young pods from proximal positions, auxin content in most inflorescences was greater than in those from the abscising distal positions. No such positional differences were observed with ABA and gibberellins. Decapitation of the plants reduced flower and pod drop from the remaining reproductive nodes. Although decapitation resulted in less abscission among distal flowers and young pods from these nodes, it did not affect the ontogenic course of auxin and ABA production in these fruits.     

Amino acid increases in fruit infested by fruit flies of the family Tephritidae

Solanum mauritianum Scop, (wild tobacco) fruit is the major host of the fruit fly Dacus cacuminatus (Hering), and is a major source of food for the brown pigeon Macropygia phasianella (Temminck) in eastern Queensland. Amino acid analyses were undertaken on fruit fly infested and uninfested S. mauritianum fruits. Infested fruits contained approximately twice the level of protein and essential amino acids compared to uninfested fruit. This increase is probably due to the plant adding additional amino acids to infested tissue and the accompanying growth of bacteria established in the fruit during oviposition. The infested fruit would provide a valuable source of protein during the pigeon breeding season.