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Ulrike Anders hat zwischen Januar und September 2005 Zähne und Gebiß rezenter Schleichkatzen (Viverridae) untersucht und Parameter identifiziert, anhand derer sich Nahrungspräferenzen zuordnen lassen. Viverriden gelten als basale Carnivoren mit omnivorem Nahrungsspektrum. Da die für echte Katzen so typische Brechschere und die Reduktion des Gebisses nur wenig ausgeprägt ist, gilt ihr Gebiss als unspezialisiert. Dennoch besitzen Viverriden Nahrungspräferenzen, die sich in der Umgestaltung ihres Gebisses, auch in einzelnen Zahnpositionen niederschlägt. Diese Veränderungen wurden metrisch charakterisiert.
1. Man findet im trockenen Monsungebiete Ost-Javas viele Baumarten, die während der Trockenzeit lange kahlstehen, nebst solchen, die während des Generalwechsels nur kurze Zeit kahl sind, und endlich eine dritte Gruppe von immergrünen Arten. Die Lauberneuerung vollzieht sich im Allgemeinen entweder mitten in der Trockenzeit oder am Ende derselben. Die Immergrünen treiben entweder immerfort an allen Knospen, soweit diese nicht in Blutenstände umgebildet werden, oder nur an einem Teil der Knospen, während bei noch anderen Arten dann und wann während einiger Zeit alle Knospen ruhen. Im Allgemeinen zeigen die jüngeren Exemplare einer beliebigen Art ein länger anhaltendes Sprosswachstnm als die älteren Vertreter. Auch in feuchtem reichem Boden findet man dieselbe Erscheinung. Im gleichmässigeren Buitenzorger Klima wird die Ruheperiode durch die in Ost-Java zeitweise kahlstehenden Arten auch wohl eingehalten, die Ruhe ist aber viel unregelmässiger und oft astweise autonom, sodass dann der Baum als Ganzes nicht kahl steht. Andere Arten stehen in Buitenzorg aber in der Trockenzeit auch während längerer Zeit kahl. Die Immergrünen sind dort auch immergrün. Einige Arten wechseln im gleichmässigeren Klima von Buitenzorg und Tjibodas alle 8 Monate das Laub und stehen dann kahl, während sie im einem Klima mit Jahresperiode einmal jährlich das Laub wechseln. 2. Es besteht ein intimer Zusammenhang zwischen Lauberneuerung und Kambialtätigkeit in dem Sinne, dass bei kahlen Bäumen das Kambium ruht und bei belaubten Bäumen, solange das Laub noch nicht zu alt .oder funktionsunfähig ist, das Kambium tätig bleibt. Bei den kahlstehenden Arten wird, soweit die Beobachtungen reichen, das Dickenwachstum durch die Laubentfaltung eingeleitet. Die in Europa häufig beobachtete Erscheinung, dass schon vor dem Knospenaufbruch das Dickenwachstum einsetzt, habe ich nie bei den kahlstehenden tropischen Arten beobachtet. Im Allgemeinen ist das Dickenwachstum desto kräftiger, je kräftiger auch das Sprosswachstum ist. 3. Zwischen der Kambialtätigkeit und der Ausbildung von Zuwachszonen tropischer Holzarten besteht die Beziehung, dass im Allgemeinen nur solche Arten scharfe ringsum geschlossene Zuwachszonen ausbilden, die zeitweise kahl stehen, also zeitweise eine Kambiumruhe aufweisen. Aber umgekehrt bilden nicht alle Arten mit periodischer Kambiumruhe auch scharfe Zuwachszonen aus. Wenn. wir ein Schema aufstellen von den logisch möglichen Beziehungen zwischen Kambialtätigkeit und Zonenbildung, dann zeigt sich dass alle Möglichkeiten auch tatsächlich in den Tropen verwirklicht sind. 4. Die Zuwachs zonen tropischer Alten werden durch eine grosse Verschiedenheit anatomischer Merkmale ausgebildet, die sich nicht nur auf die verschiedenen Arten beschränkt, sondern die man auch innerhalb derselben Art antrifft. Diese Merkmale sind: a. Eine radiale Verkürzung der letzten Spätholz-Libriformfasern. b. Ein rund herum laufender schmaler Parenchymstreifen. Sehr oft werden diese Parenchymzellen als letzte Elemente des Dickenwachstums vor der Ruhe ausgebildet. Bisweilen entsteht eine Zone von dünnwandigem Parenchym aus der Zwischenzone von noch nicht ausgewachsenen Zellen zwischen Kambium und Altholz, die während der Ruhe unverändert liegen bleiben. c. Ein schmaler Streifen Libriform ohne Gefässe oder Parenchym, oft als erste Bildung des Jungholzes. d. Eine Periodizität in der Breite der abwechselnden Libriform- und Parenchymbänder. e. Eine Periodizität in der Gefässgrässe oder der Gefässanordnung. Oft findet man im Frühholz eine Anhäufung grösserer Gefässe, ziemlich oft sind aber au.ch die Gefässe in der Mitte der Zuwachszone am grössten; bisweilen findet man eine Anhäufung kleinerer Gefässe im Spätholz, während das Frühholz dann wohl grössere Gefässe aufweist, die aber weiter voneinander entfernt sind. Bisweilen ist auch das Frühholz ausgesprochen gefässarm. 5. Es gibt nur verhältnissmässig wenige tropische Holzarten, bei denen die Ausbildung der Zuwachszonen unter gleichmässigen äusseren Umständen ganz ausbleibt, während bei ungleichmässi gen Aussenfaktoren schöne Zuwachszonen ausgebildet werden. Die anderen Arten bilden entweder gleich deutliche Zuwachszonen in Ost- und West-Java aus (es sind dies die Arten mit fester Laubperiodizität) oder die Zonen sind im gleichmässigeren Klima von West-Java etwas weniger, scharf oder etwas unregelmässiger als in Ost-Java. Die Ausbildung von Zuwachszonen bleibt sehr oft in der Jugend aus, oder die in den ersten Lebensjahren gebildeten Zonen sind unvollkommen und verwaschen. 6. Die Altersbestimmung durch Abzählung der Zuwachszonen ist in den Tropen nicht so zuverlässig wie in den gemässigten Gegenden, weil oft die Zuwachszonen etwas unregelmftssig sind und dabei die Zuwachszonen in der Jugend bisweilen fehlen. Es gibt aber eine Anzahl Holzarten, die im periodischen Klima von Ost-Java bis auf einige Jahre genaue Resultate geben, während bei anderen die Fehlergrenze innerhalb 20-30% bleibt. In West-Java ist die Abzählung der Zuwachszonen für viele dieser Arten nicht mehr zuverlässig; bei einigen Arten wird man jedoch auch dort bis auf einige Jahren genaue Resultate bekommen, bei anderen bis auf 20-30% genaue Resultate. 7. Die aus kälteren Gegenden in Tjibodas eingeführten Holzarten sind dort grösstenteils immergrün; die Lau bhö1zer jedoch zeigen meistens eine astweise Periodizität. Alnus, Rosa und Sambuctts treiben immerfort an allen unverletzten Knospen. Das Dickenwachstum steht in demselben Zusammenhang mit der Laubperiodizität, wie wir es für die tropischen Arten gefunden haben. Die meisten Laubholzarten mit astweiser Laubperiodizität bilden ganz normale Zuwachszonen aus, die meistens der Zahl der Trieb ab sätze entsprechen. In den dickeren Ästen und im Hauptstamm findet man dann aber bisweilen Unregelmässigkeiten, wie zusammenfliessende oder blind im Gewebe endigende Zonen. Die immer durchwachsenden Laubholzarten zeigten oft keine oder nur vage Zuwachszonen. Die Koniferen zeigen dort sehr oft unscharfe Zuwachszonen, bei denen das Spätholz nach beiden Seiten allmählig in Frühholz übergeht. 8. Durch Rindenringelungs-, Entknospungs- und Verdunkelungsversuche an laubverlierenden und immergrünen tropischen Holzarien, sowie aus einer kritischen Übersicht der wichtigeren Literatur, ist festgestellt, dass das erneute Dickenwachstum nach der Ruhe durch in der Rinde weiter transportierte Stoffe oder Reize angeregt wird, die von den sich entfaltenden Organen ausgehen. Wenn die austreibenden Organen entfernt oder durch eine Rindenringelung isoliert werden, bildet sich kein oder nur sehr wenig neues Holz aus. Versuche haben festgestellt, dass auch bei Bäumen in Europa, die schon Kambialtätigkeit zeigen bevor noch die Knospen aufgebrochen sind, der Reiz zum Dickenwachstum von den Knospen ausgeht (S. 77/8). Die Unterbrechung des absteigenden Saftstromes bei Immergrünen oder bei belaubten Exemplaren laubverlierender Arten, hat ein sofortiges oder sehr baldiges Einstellen des Dickenwachstums zufolge, das aber wieder aufgenommen wird, nachdem der Saftstrom wieder ungehindert durchgeht. Das Dickenwachstum tritt auch bei austreibenden verdunkelten Pflanzen auf, deren Assimilationstätigkeit verhindert ist. Bei den Immergrünen sind es hauptsächlich die auswachsenden Organe und weniger die alten Blattmassen, die den Reiz zum Dickenwachstum abgeben. Das erneute Dickenwachstum bei in kahlem Zustande blühenden Arten wird schon durch die auswachsenden Blüten angeregt. 9. Bei der plötzlichen Unterbrechung des absteigenden Saftstromes wird das Dickenwachstum, entweder sofort oder sehr bald, ganz eingestellt und bleibt die Zone halb ausgewachsener Zellen zwischen Kambium und fertigem Holze entweder unverändert liegen oder es wachsen die Gefässanlagen zu kleinen Gefässen mit paratrachealem Parenchym aus, während das zwischenliegende Gewebe parenchymatisch bleibt. Nach erneutem Dickenwachstum wächst diese Zone dann meistens nicht weiter aus. Daher weisen bei plötzlicher Unterbrechung des absteigenden Saftstromes (z. B. bei Rindenringelung oder Entblätterung), die dadurch hervorgerufenen Zuwachszonen meistens, je nach der Wuchskraft, eine schmälere oder breitere Zone dünnwandigeren Gewebes auf. 10. Aus verschiedenen Versuchen und Beobachtungen geht hervor, dass die jungen sich entfaltenden Organe die Bildung des weitlumigen Frühholzes verursachen, während das englumigere Spätholz durch die Tätigkeit der älteren, schon ausgewachsenen Blätter hervorgerufen wird. Hieraus ergeben sich die Bedingungen zur künstlichen Herstellung von Zuwachszonen. Vor allem muss die Ausbildung des Spätholzes schon eingetreten sein. Wenn das nicht der Fall ist, kann sie künstlich durch das während einiger Zeit fortgesetzte Abschneiden der Sprosspitze und der treibenden Knospen hervorgerufen werden. Dann muss eine Zone Jungholz ausgebildet werden, indem ein kräftiges Austreiben der Knospen herbeigeführt wird. Solche künstliche Zuwachszonen wurden tatsächlich hergestellt. 11. Es wird die schon vorher durch andere Verfasser aufgestellte Hypothese verteidigt, dass das Dickenwachstum von der Laubtätigkeit verursacht wird, entweder durch Ausbildung von Hormonen oder durch einen in der Rinde herabbeförderten Reiz. Die Bildung von Hormonen ist wahrscheinlicher als Reizwirkung. Auch die Ausbildung von Zuwachszonen wird dieser Ursache zugeschrieben; Die von anderen Autoren angeführten Faktoren, die das Dickenwachstum und die Ausbildung von Zuwachszonen hervorrufen sollen, wie Rindenspannung, Turgor der Jungholzzellen, Ernährung des Kambiums mit Nährsalzen und organischen Nährstoffen, Wassergehalt der Tungholzregion, haben entweder nur eine indirekte Wirkung, indem sie das Laubwachstum beeinflussen, oder sie werden selbst durch die vom Laube ausgehenden Stoffe oder Reize hervorgerufen. Ein gewisser Zusammenhang zwischen Transpiration und Ausbildung von Wasserleitungsbahnen ist in vielen Fällen auch nicht zu verkennen, aber dieser Zusammenhang ist wahrscheinlich nur indirect und wird direct durch vorn Laube ausgehende Reizstoffe oder Reize hervorgerufen. Die Frage des Dickenwachstums und der Jahresring (Zuwachszonen) bildung wird also zurückgeführt auf eine andere Erscheinung im Pflanzenleben, die Laubperiodizität ; dieser letzte Satz aber mit der Einschränkung, dass in erster Instanz die erbliche Anlage darüber entscheidet, ob überhaupt, und in wie weit, Zuwachszonen ausgebildet werden können. 12. Es wird betont dass man keine palaöklimatologische Schlüsse aus vereinzelten Beobachtungen ziehen darf; wenn man aber bei vielen Arten scharfen, ringsum geschlossenen Zuwachszonen antrifft, so darf man daraus wohl auf ein periodisch kälteres Klima schliessen.
Lauberneuerung und andere periodische Lebensprozesse in dem trockenen Monsungebiet Ost-Java's
(1923)
Hasselmann, Martin ; Vekemans, Xavier ; Pflugfelder, Jochen ; Koeniger, Nikolaus ; Koeniger, Gudrun ; Tingek, Salim ; Beye, Martin
Our understanding of the impact of recombination, mutation, genetic drift and selection on the evolution of a single gene is still limited. Here we investigate the impact of all of these evolutionary forces at the complementary sex determiner (csd) gene which evolves under a balancing mode of selection. Females are heterozygous at the csd gene and males are hemizygous; diploid males are lethal and occur when csd is homozygous. Rare alleles thus have a selective advantage, are seldom lost by the effect of genetic drift and are maintained over extended periods of time when compared to neutral polymorphisms. Here, we report on the analysis of 17, 19 and 15 csd alleles of Apis cerana, Apis dorsata and Apis mellifera honey bees respectively. We observed great heterogeneity of synonymous (pi S) and nonsynonymous (pi N) polymorphisms across the gene, with a consistent peak in exon 6 and 7. We propose that exons 6 and 7 encode the potential specifying domain (csd-PSD) which has accumulated elevated nucleotide polymorphisms over time by balancing selection. We observed no direct evidence that balancing selection favors the accumulation of nonsynonymous changes at csd-PSD (pi N/pi S ratios are all < 1, ranging from 0.6 to 0.95). We observed an excess of shared nonsynonymous changes, which suggests that strong evolutionary constraints are operating at csd-PSD resulting in the independent accumulation of the same nonsynonymous changes in different alleles across species (convergent evolution). Analysis of a csd-PSD genealogy revealed relatively short average coalescence times (~6 million years), low average synonymous nucleotide diversity (pi S < 0.09) and a lack of trans-specific alleles which substantially contrasts with previously analyzed loci under strong balancing selection. We excluded the possibility of a burst of diversification after population bottlenecking and intragenic recombination as explanatory factors, leaving high turn-over rates as the explanation for this observation. By comparing observed allele richness and average coalescence times with a simplified model of csd-coalescence, we found that small long term population sizes (i.e. Ne <104), but not high mutation rates, can explain short maintenance times, implicating a strong impact of genetic drift on the molecular evolution of highly social honey bees.
Molina, Carlos ; Rotter, Björn ; Horres, Ralf ; Udupa, Sripada M. ; Besser, Bert ; Bellarmino, Luis ; Baum, Michael ; Matsumura, Hideo ; Terauchi, Ryohei ; Kahl, Günter ; Winter, Peter
Background Drought is the major constraint to increase yield in chickpea (Cicer arietinum). Improving drought tolerance is therefore of outmost importance for breeding. However, the complexity of the trait allowed only marginal progress. A solution to the current stagnation is expected from innovative molecular tools such as transcriptome analyses providing insight into stress-related gene activity, which combined with molecular markers and expression (e)QTL mapping, may accelerate knowledge-based breeding. SuperSAGE, an improved version of the serial analysis of gene expression (SAGE) technique, generating genome-wide, high-quality transcription profiles from any eukaryote, has been employed in the present study. The method produces 26 bp long fragments (26 bp tags) from defined positions in cDNAs, providing sufficient sequence information to unambiguously characterize the mRNAs. Further, SuperSAGE tags may be immediately used to produce microarrays and probes for real-time-PCR, thereby overcoming the lack of genomic tools in non-model organisms. Results We applied SuperSAGE to the analysis of gene expression in chickpea roots in response to drought. To this end, we sequenced 80,238 26 bp tags representing 17,493 unique transcripts (UniTags) from drought-stressed and non-stressed control roots. A total of 7,532 (43%) UniTags were more than 2.7-fold differentially expressed, and 880 (5.0%) were regulated more than 8-fold upon stress. Their large size enabled the unambiguous annotation of 3,858 (22%) UniTags to genes or proteins in public data bases and thus to stress-response processes. We designed a microarray carrying 3,000 of these 26 bp tags. The chip data confirmed 79% of the tag-based results, whereas RT-PCR confirmed the SuperSAGE data in all cases. Conclusion This study represents the most comprehensive analysis of the drought-response transcriptome of chickpea available to date. It demonstrates that – inter alias – signal transduction, transcription regulation, osmolyte accumulation, and ROS scavenging undergo strong transcriptional remodelling in chickpea roots already 6 h after drought stress. Certain transcript isoforms characterizing these processes are potential targets for breeding for drought tolerance. We demonstrate that these can be easily accessed by micro-arrays and RT-PCR assays readily produced downstream of SuperSAGE. Our study proves that SuperSAGE owns potential for molecular breeding also in non-model crops.