几十年来,人们一直认为韧皮部筛元素(SE)闭塞是抵抗韧皮部汁液喂养昆虫的一种机制。尽管这很可能是一种普遍现象,但很少有研究检验了这一假设。这篇综述集中在call和P蛋白的SE闭塞。两者都是可逆的,这将使植物在SEs被穿透时能够防御韧皮部汁液喂食器,并在昆虫放弃并撤回其探针时恢复正常功能。callose(β-1,3葡聚糖与一些β-1,6分支)在许多不同组织的植物生理学中起着许多作用,每个都在不同call糖合酶基因的控制下;只有沉积在SE筛孔中的call糖才与SE闭塞有关。筛孔中call糖的量(以及它阻碍树液流动的程度)取决于call糖合酶和β-1,3葡聚糖酶之间的活性平衡。在一些研究中,筛孔call质沉积已被证明对某些韧皮部树液饲养者具有抵抗力,在一个,易感和抗性水稻品种之间的抗性差异是由于昆虫能够或无法上调植物β-1,3葡聚糖酶降解call的沉积。P蛋白仅存在于双子叶植物中,包括多种蛋白质,并非所有这些都涉及SE闭塞。在一些植物中,P蛋白在成熟的功能性SE中形成不同的体。在乳头状豆科植物中,这些离散的身体,称为forisomes可以扩张和收缩。在他们的扩张状态下,它们在收缩状态下有效地堵塞SE并阻止SAP的流动,它们对汁液流动的阻力可以忽略不计。孔体的扩张是由Ca2流入SE引发的。不适应豆科植物的通才蚜虫对豆科植物(Viciafaba)SE的渗透会引发毒体扩张,从而阻塞SE并阻止蚜虫摄取汁液。相比之下,一个豆类专家蚜虫,Acyrthosiphonpisum,不会引发有害的扩张,并且很容易从V.faba中摄取汁液。非豆类SE中的P蛋白体似乎不参与SE闭塞。在大多数双子叶植物中,P蛋白不形成离散体,而是以丝状聚集体的形式出现,粘附在SE的顶叶边缘,并响应损伤,被释放到内腔中,在那里它们被汁液流带到下游筛板,在那里它们回流并堵塞筛孔。它们在实际阻止汁液流动方面的有效性是有争议的。在一项研究中,它们似乎对汁液的流动几乎没有阻力,而在其他研究中,他们提供了相当大的阻力。为了应对甜瓜的伤害,它们完全阻止了树液的流动,在一个抗蚜虫的甜瓜里,甜瓜蚜虫穿透SEs,山雀,引发P蛋白闭塞,阻止蚜虫摄取汁液。第一个P蛋白被描述,PP1,仅发生在南瓜属,尽管它经常被认为是一种SE闭塞蛋白,实验证据表明,它在SE闭塞中没有重要作用。韧皮部汁液饲养者减轻P蛋白闭塞的最常见策略似乎是避免触发它。一项广泛引用的体外研究表明,蚜虫唾液可以逆转P蛋白闭塞,但随后的一项研究表明,唾液在体内逆转P蛋白闭塞方面无效。最后,据推测,由俄罗斯小麦蚜虫引发的小麦中的SEcall质沉积会产生有利于蚜虫的人工水槽,但需要更多的研究来检验这一假设。
Phloem sieve element (SE) occlusion has been hypothesized for decades to be a mechanism of resistance against phloem sap-feeding insects. Few studies have tested this hypothesis although it is likely a widespread phenomenon. This
review focuses on SE occlusion by callose and P-proteins. Both are reversible, which would allow the plant to defend itself against phloem sap-feeders when SEs are penetrated and resume normal function when the insects give up and withdraw their stylets. Callose (β-1,3 glucans with some β-1,6 branches) serves many roles in plant physiology in many different tissues, each being under the control of different callose synthase genes; only callose deposited in SE sieve pores is relevant to SE occlusion. The amount of callose in sieve pores (and consequently how much it impedes sap flow) is determined by the balance in activity between callose synthase and β-1,3 glucanase. Sieve pore callose deposition has been shown to provide resistance to some phloem sap-feeders in a few studies, and in one, the difference in resistance between a susceptible and resistant rice variety was due to the ability or inability of the insect to upregulate the plants\' β-1,3 glucanase that degrades the callose deposition. P-proteins occur only in dicotyledons and include a variety of proteins, not all of which are involved in SE occlusion. In some plants, P-proteins form distinct bodies in mature functional SEs. In papilionid legumes, these discrete bodies, called forisomes can expand and contract. In their expanded state, they effectively plug SEs and stop the flow of sap while in their contracted state, they provide negligible resistance to sap flow. Expansion of forisomes is triggered by an influx of Ca2+ into the SE. Penetration of a legume (Vicia faba) SE by a generalist aphid not adapted to legumes triggers forisome expansion which occludes the SE and prevents the aphid from ingesting sap. In contrast, a legume specialist aphid, Acyrthosiphon pisum, does not trigger forisome expansion and readily ingests sap from V. faba. P-protein bodies in SEs of non-legumes do not appear to be involved in SE occlusion. In most dicotyledons, P-proteins do not form discrete bodies, but rather occur as filamentous aggregations adhering to the parietal margins of the SE and in response to damage, are released into the lumen where they are carried by the flow of sap to the downstream sieve plate where they back up and clog the sieve pores. Their effectiveness at actually stopping the flow of sap is controversial. In one study, they seemed to provide little resistance to the flow of sap while in other studies, they provided considerable resistance. In response to injury in melon, they completely stop the flow of sap, and in an aphid-resistant melon, penetration of SEs by the melon aphid, Aphis gossypii, triggers P-protein occlusion which prevents the aphids from ingesting sap. The first P-protein described, PP1, occurs only in the genus Cucurbita, and although it has been often cited to function as a SE occlusion protein, experimental evidence suggests it does not play a significant role in SE occlusion. The most common strategy for phloem sap-feeders to mitigate P-protein occlusion seems to be avoid triggering it. A widely cited in vitro study suggested that aphid saliva can reverse P-protein occlusion, but a subsequent study demonstrated that saliva was ineffective at reversing P-protein occlusion in vivo. Lastly, SE callose deposition in wheat triggered by Russian wheat aphid has been hypothesized to create an artificial sink that benefits the aphid, but additional studies are needed to test that hypothesis.