Premise of research. In Caryophyllaceae with a gynoecium isomerous with the other floral whorls, carpels can be in either antesepalous or antepetalous position. Morphologists have long been unable to find a unifying hypothesis to explain the different carpel positions in Caryophyllaceae, as variation in carpel position breaks with Hofmeister’s rule of alternating whorls. Methodology. In this study, we investigate the floral development of six species of Caryophyllaceae with isomerous flowers using scanning electron microscopy to clarify the differences between carpel positions and whether developmental modes are responsible. Special attention is paid to the correlation between the development of carpels and other floral organs. Pivotal results. In Spergula arvensis, Agrostemma githago, and Myosoton aquaticum, carpels are in antepetalous position, while the antesepalous stamens develop more centrally in a sequence inverse to the sepals. In Cerastium glomeratum and Silene diocia, carpels are in antesepalous position, while the antesepalous stamens do not show an inverse development and are inserted almost at the same level as antepetalous stamens. In Lychnis flos-cuculi, the carpels are not clearly in an antesepalous or antepetalous radius, and the development of antesepalous stamens shows a slight unequal growth. Conclusions. Developmental evidence rejects the hypothesis that the developmental modes of carpels are the reason for the different carpel positions. Carpel position is correlated with the developmental sequence of antesepalous stamens and the level of emergence of antepetalous stamens on the floral apex. Space availability and pressure exercised by the sepals is responsible for the unequal development and a more central position of antesepalous stamens, with those opposite the inner sepals developing faster, causing the carpels to shift to an antepetalous position. With greater availability of space, antesepalous stamens initiate simultaneously or continue the sequence of sepal initiation, and all stamens are more or less equidistant from the apex; the antesepalous stamens do not display an inverse development, and the carpels initiate in antesepalous position, which is consistent with Hofmeister’s rule.

中文翻译：

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霍夫迈斯特法则悖论：解释石竹科多变的心皮位置

研究的前提。在具有雌蕊与其他花轮同分异构体的石竹科中，心皮可以处于对瓣或对瓣位置。长期以来，形态学家一直无法找到一个统一的假设来解释石竹科中不同的心皮位置，因为心皮位置的变化违反了霍夫迈斯特的轮换规则。方法。在这项研究中，我们使用扫描电子显微镜研究了六种具有同分异构花的石竹科植物的花发育，以阐明心皮位置之间的差异以及发育模式是否是负责任的。特别注意心皮与其他花器官发育之间的相关性。关键结果。在 Spergula arvensis、Agrostemma githago 和 Myosotonaquaticum 中，心皮位于前瓣位置，而对萼雄蕊以与萼片相反的顺序发育更集中。在 Cerastium glomeratum 和 Silene diocia 中，心皮处于对瓣位置，而对瓣雄蕊不显示反向发育，几乎与对瓣雄​​蕊处于同一水平。在Lychnis flos-cuculi中，心皮在antesepalous或antepetalous半径内不明显，并且antesepalous雄蕊的发育显示出轻微的不均匀生长。结论。发育证据否定了心皮的发育模式是导致不同心皮位置的原因的假设。心皮位置与对瓣雄蕊的发育顺序和花尖上对瓣雄蕊的出现水平有关。萼片的空间可用性和压力是造成对萼雄蕊发育不均和更中心位置的原因，与内萼片相对的雄蕊发育更快，导致心皮转移到前瓣位置。随着空间的更大可用性，对萼雄蕊同时开始或继续萼片开始的顺序，并且所有雄蕊或多或少与先端等距；对生雄蕊不显示反向发育，心皮起始于对生位置，这与 Hofmeister 规则一致。对萼雄蕊同时起始或延续萼片起始序列，所有雄蕊或多或少与先端等距；对生雄蕊不显示反向发育，心皮起始于对生位置，这与 Hofmeister 规则一致。对萼雄蕊同时起始或延续萼片起始序列，所有雄蕊与先端或多或少等距；对生雄蕊不显示反向发育，心皮起始于对生位置，这与 Hofmeister 规则一致。