多功能葉片測(cè)量?jī)x助力研究胚胎受損種子對(duì)植株再生的貢獻(xiàn)

摘要 

種子捕食中的胚胎損傷是一種常見(jiàn)現(xiàn)象，通常被認(rèn)為相當(dāng)于種子死亡。我們假設(shè)胚胎（胚根加胚芽）比例較大的種子對(duì)嚙齒類(lèi)動(dòng)物的種子損傷具有更強(qiáng)的耐受性，從而使種子能夠成功萌發(fā)。為了驗(yàn)證這一假設(shè)，我們通過(guò)比較完整和胚胎受損種子的傳播模式以及它們?cè)趦煞N胚胎大小不同的橡樹(shù)中出苗的生態(tài)生理反應(yīng)，檢查了發(fā)芽情況，并估計(jì)了胚胎受損種子對(duì)植株再生的貢獻(xiàn)。我們的結(jié)果表明，兩種橡樹(shù)的胚胎大小與種子大小呈正相關(guān)，但其中一種（櫟屬）的胚胎比另一種（農(nóng)業(yè)櫟屬）的胚胎長(zhǎng)，表明胚胎大小存在種間差異。胚胎切除行為的概率（部分種子損傷以去除胚胎）與種子大小呈正相關(guān)，但所有大小的損傷強(qiáng)度相對(duì)恒定。胚胎比例較長(zhǎng)的物種表現(xiàn)出更強(qiáng)的從胚胎損傷中再生的能力，因?yàn)檩^長(zhǎng)的胚胎經(jīng)歷嚙齒類(lèi)動(dòng)物對(duì)胚胎的總（致命）損傷的概率降低。種子大小和種內(nèi)胚胎大小只會(huì)增加胚胎較短的物種的發(fā)芽成功率，從而使較大的種子避免完全的胚胎損傷。與胚胎受損種子相比，完整種子的幼苗表現(xiàn)更好（更大的植物生物量和更高的葉綠素和氮指數(shù)）。然而，胚胎受損種子的幼苗顯示出較高的花青素含量，這可能是對(duì)種子受損的一種反應(yīng)。完整和胚胎受損的橡子在嚙齒類(lèi)動(dòng)物身上表現(xiàn)出相似的傳播模式（例如種子貯藏行為和傳播距離）。重要的是，對(duì)于胚胎受損的種子，嚙齒類(lèi)動(dòng)物的種子回收率明顯較低，使種子存活率更高，這支持胚胎切除行為是一種長(zhǎng)期儲(chǔ)存種子的策略的預(yù)期。我們的結(jié)論是，對(duì)胚胎損傷的耐受性是一種重要的繁殖策略，可以促進(jìn)植物的繁殖。胚胎大小在種子對(duì)動(dòng)物捕食的進(jìn)化和生態(tài)反應(yīng)中起著至關(guān)重要的作用。 

關(guān)鍵詞：胚胎切除，發(fā)芽成功，嚙齒動(dòng)物種子捕食，種子傳播，幼苗生理學(xué)，種子大小 
Embryo size as a tolerance trait against seed predation: Contribution of embryo-damaged seeds to plant regeneration 

Abstract 
Embryo damage in seed predation is a common occurrence and has been generally considered equivalent to seed death. We hypothesize that seeds with proportionally larger embryos (radicle plus plumule) provide greater tolerance to seed damage by rodents, allowing successful germination. To test this hypothesis, we examined germination and estimated the contribution of embryo-damaged seeds to plant regeneration by comparing the dispersal patterns of intact and embryo-damaged seeds and the ecophysiological responses of their emerged seedlings in two oak species of contrasting embryo size. Our results show that embryo size was positively correlated with seed size in both oak species, but one (Quercus lobata) had proportionally longer embryos than the other (Q. agrifolia), revealing inter-specific differences in embryo size. Probability of embryo excision behavior (partial seed damage to extirpate the embryo) was positively associated with seed size but intensity of damage was relatively constant across all sizes. The species with proportionally longer embryos showed a stronger capacity to regenerate from embryo damage because longer embryos experienced reduced probability of total (lethal) embryo damage by rodents. Seed size and thus intraspecific embryo size only increased germination success in the species with shorter embryos, allowing larger seeds to escape full embryo damage. Seedlings from intact seeds performed better (greater plant biomass and higher chlorophyll and nitrogen index) than those from embryo-damaged seeds. However, seedlings from embryo-damaged seeds showed higher anthocyanin content, a possible response to seed damage. Intact and embryo-damaged acorns showed similar dispersal patterns by rodents (e.g. seed caching behavior and dispersal distances). Importantly, seed retrieval by rodents was significantly lower for embryo-damaged seeds, allowing greater seed survival and supporting the expectation that embryo excision behavior is a strategy to store seeds for longer periods. We conclude that tolerance to embryo damage is an important reproductive strategy that enables plant recruitment. Embryo size could play a crucial role in the evolutionary and ecological responses of seeds to animal predation.

Keywords
Embryo excision
Germination success
Rodent seed predation
Seed dispersal
Seedling physiology
Seed size
 

Dualex是一款源自于法國(guó)國(guó)家科學(xué)院 (CNRS)及巴黎第十一大學(xué)技術(shù)，由奧地利PESSL公司生產(chǎn)（原法國(guó)Force-A公司）開(kāi)發(fā)的新型多功能葉片測(cè)量?jī)x。它可同時(shí)準(zhǔn)確測(cè)量葉片的葉綠素含量、葉片表層的類(lèi)黃酮和花青素含量，適用于植物生理學(xué)和農(nóng)學(xué)（如水稻葉綠素濃度，玉米氮素狀況，葡萄藤等）相關(guān)研究。其測(cè)量對(duì)象可以是單子葉植物，雙子葉植物或多年生植物。這款設(shè)備簡(jiǎn)單易用，可進(jìn)a行實(shí)時(shí)和非破壞性測(cè)量。由于不需要校準(zhǔn)標(biāo)定和事先的樣品制備，測(cè)量工作可在實(shí)驗(yàn)室或現(xiàn)場(chǎng)完成。此外，該設(shè)備在各種溫度的和環(huán)境光照條件下均可正常使用。系統(tǒng)采用專(zhuān)利設(shè)計(jì)，光學(xué)傳感器可進(jìn)行簡(jiǎn)單、快速、無(wú)損測(cè)量葉片中的葉綠素、多酚以及花青素。

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