Summary
This paper deals with the influence of environmental factors, particularly those that may be expected to change the rate of growth in the larval period, on the homoeotic wing-like outgrowths in the eyes of the strainloboid-ophthalmoptera (see Ouweneel, 1969a, b).
The penetrance of the homoeotic effect increases with maternal age (Table 1); Delcour (1968) furnished evidence that there is a relation between maternal age and growth rate of tissues in the larva. Penetrance is inversely related to temperature; at low temperatures the development of the whole larva is strongly retarded, but the growth of the eye disc is probably less retarded, so that it grows relatively more than at higher temperatures. It is notable that the temperature-sensitive period (24–60 h after hatching, at 25° C; Fig. 1) precedes, and coincides with, the period of abnormal hyperplasia in the disc (Ouweneel, 1969b).
Acetamide, given in the food, enhances the number of facets so that even the wild type eye size may be reached; in proportion to its concentration it moreover strongly increases the penetrance of the wing-like outgrowths. At higher concentrations, however, when eye enlargement is maximal, the penetrance of the outgrowths decreases again (Figs. 2–5). The optimal concentration (at which the penetrance is maximal) is higher at higher temperatures (Figs. 2–5; cf. Fig. 7). The sensitive period for acetamide extends from about 0–55 h after hatching (at 25° C) (Fig. 6). Uracil increases the penetrance of the homoeotic effect (Table 2). Sodium tetraborate seems to increase the penetrance of the homoeotic outgrowths, but to reduce their expressivity slightly at higher concentrations. Starvation (Tabel 3) and farnesol medium prolong the larval period, but do not show clear-cut effects on the homoeotic phenomenon, probably because they only postpone metamorphosis, and do not retard the whole of larval development (compare temperature effect, above). UV irradiation influenced penetrance, but lower dosages had a greater effect than higher ones; the former suppressed the homoeotic effect at early larval ages, but stimulated it at later ages; this points to the occurrence of two separate sensitive periods for UV (Table 4). In all experiments clear-cut sexual differences in sensitivity were observed, which may be ascribed to the sex-linked location of theophthalmoptera modifier (Ouweneel, 1969a).
Evidence can be found in the literature that enhanced proliferation is a prerequisite for many homoeotic phenomena in the broad sence (cf. Fig. 8). The present study and previous data show that also homoeotic mutations often interact with agents causing changes in growth rate; it is possible that the basic action of all homoeotic mutations has to do with such changes.
Zusammenfassung
Es wurde der Einfluß wachstumsändernder Umweltfaktoren während der Larvalentwicklung auf die homöotischen, flügelähnlichen Auswüchse im Auge des Stammesloboid-ophthalmoptera untersucht.
Die Penetranz des homöotischen Effekts steigt an mit dem Alter der Mutterfliegen; nach Literaturangaben besteht ein Zusammenhang zwischen mütterliches Alter und Wachstumsgeschwindigkeit der Gewebe während der Larvalentwicklung. Zwischen Penetranz und Temperatur besteht ein umgekehrtes Verhältnis; dieses wird erklärt durch die Annahme, daß obwohl die Entwicklung der ganzen Larve bei niedriger Temperatur stark verzögert wird, das Wachstum der Augenimaginalscheibe weniger stark gehemmt wird, so daß die Augenscheibe relativ mehr wächst als bei höheren Temperaturen. Die temperaturempfindliche Periode (24–60 Std nach dem Schlüpfen bei 25° C) bestreicht die Periode der erhöhten Hyperplasie in der Augenscheibe (Ouweneel, 1969b).
Azetamid im Futter erhöht die Zahl der Augenfazetten, manchmal bis zur normalen Augengröße; daneben erhöht es die Penetranz des homöotischen Effekts beträchtlich in Abhängigkeit der Konzentration. Allerdings sinkt die Penetranz bei Konzentrationen, die eine maximale Augenvergrößerung bewirken, wieder ab. Die optimale Konzentration (bei der die Penetranz maximal ist) ist höher, je höher die Temperatur. Die empfindliche Periode für Azetamid erstreckt sich von etwa 0 bis 55 Std nach dem Schlüpfen (bei 25° C). Urazil erhöht die Penetranz des homöotischen Effekts. Natriumtetraborat erhöht die Penetranz, setzt aber die Expressivität bei höheren Konzentrationen leicht herab.
Hüngern und Farnesolgaben im Futter verlängern die Larvalentwicklung, ohne jedoch den homöotischen Effekt eindeutig zu beeinflussen, wahrscheinlich weil sie nur die Metamorphose hinausschieben, ohne die gesamte Larvalentwicklung zu verzögern (siehe oben bei Temperatureffekt). Ultraviolettbestrahlung hat in niedrigen Dosierungen einen stärkeren Effekt auf die Penetranz als in hohen Dosierungen. Bei jungen Larven wird die Penetranz herabgesetzt, bei älteren Larven dagegen erhöht, was auf zwei unterschiedene empfindliche Perioden für Ultraviolett hinweist. In allen Experimenten wurden deutliche Geschlechtsunterschiede in der Empfindlichkeit festgestellt, die der geschlechtsgebundenen Lokalisation desophthalmoptera-Modifikators zugeschrieben werden können.
Aus Literaturangaben läßt sich schliessen, daß erhöhte Proliferation eine Voraussetzung ist für viele homöotische Erscheinungen im weitesten Sinne (einschließlich Transdetermination). Es wird auf die Möglichkeit hingewiesen, daß alle homöotischen Mutationen sich grundsätzlich durch Änderungen der Wachstumsgeschwindigkeit auswirken.
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Aided by a grant from the Netherlands Organization for the Advancement of Pure Research (Z.W.O.).
I am much indebted to Dr. J. Faber for stimulating discussions and for critically reading the manuscript. I thank Mrs. M. van Deventer for accurately carrying out many of the experiments. Prof. P. D. Nieuwkoop kindly read the manuscript and suggested several improvements.
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Ouweneel, W.J. Influence of environmental factors on the homoeotic effect ofloboid-ophthalmoptera inDrosophila melanogaster . W. Roux' Archiv f. Entwicklungsmechanik 164, 15–36 (1969). https://doi.org/10.1007/BF00577679
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DOI: https://doi.org/10.1007/BF00577679