跟著城市嚮導「老臺北胃」，用味道認識臺北

很多朋友來臺北，
都會問我同一個問題：
「臺北小吃那麼多，到底該從哪裡開始吃？」
夜市裡攤位一字排開、老店藏在巷弄轉角，
看起來都很有名，卻又怕吃錯、踩雷，
結果行程走完，反而沒真正記住臺北的味道。
我常被朋友笑說是「老臺北胃」。
不是因為特別會吃，而是因為在這座城市待久了，
知道哪些味道是陪著臺北人成長的日常。
這篇文章，就是我整理的一份清單。
如果你第一次來臺北，
我會帶你從這 10 樣最具代表性的臺北小吃開始，
不追一時爆紅、不走浮誇路線，
而是讓你吃完後能真正理解
原來，這就是臺灣的小吃文化。
跟著老臺北胃走，
用最簡單的方式，
把臺北的味道，一樣一樣記在心裡。

我怎麼選出這 10 大臺北小吃？

在臺北，
你隨便走進一條夜市或老街，
都可以輕易列出 30 種以上的小吃。
所以這份清單，
不是「臺北最好吃」的排名，
 而是我站在「第一次來臺北的旅客」角度，
做的推薦。
身為一個被朋友稱作「老臺北胃」的人，
我選這 10 樣小吃時，心裡一直放著幾個原則。

一吃就知道：這就是臺灣味

燒烤、火鍋很好吃，
但換個城市、換個國家，也吃得到。
我挑的，是那種
只要一入口，就會讓人聯想到的臺灣味。
 不需要解釋太多，舌頭就能懂。

不只是好吃，而是有「臺北日常感」

臺北的小吃迷人，
不只在味道，
而在它融入生活的方式。
我在意的是：

1. 會不會出現在早餐、宵夜、下班後
2. 有沒有陪伴這座城市很久的記憶

吃完之後，你會記得臺北

最後一個標準很簡單。
如果你回到家，
還會突然想起某個味道、某碗熱湯、某個攤位的香氣
那它就值得被放進這份清單裡。

接下來的 10 樣臺北小吃，
就是我會親自帶朋友去吃的在地美食。
不趕行程、不拚數量，
而是一口一口，
慢慢認識臺北。

第 1 家：饌堂－黑金滷肉飯（雙連店）｜一碗就懂臺灣人的日常

如果只能用一道料理，
 來解釋臺灣人的日常飲食，
 那我一定會先帶你吃滷肉飯。
在臺北，滷肉飯不是什麼特別的節慶料理，
 而是從早餐、午餐到宵夜，
 默默陪著很多人長大的味道。
而在眾多滷肉飯之中，
饌堂－黑金滷肉飯（雙連店），
 我很常帶第一次來臺北的朋友造訪的一家。

為什麼第一站，我會選饌堂？
饌堂的滷肉飯，走的是**「黑金系」路線**。
滷汁顏色深、香氣厚，
卻不死鹹、不油膩。
滷肉切得細緻，
肥肉入口即化，搭配熱騰騰的白飯，
每一口都是很完整、很臺灣的味道。
對第一次吃滷肉飯的旅客來說，
這種風味夠經典、也夠穩定，
不需要太多心理準備，就能理解為什麼臺灣人這麼愛它。

不只是好吃，而是「現在的臺北感」
饌堂並不是那種躲在深巷裡的老攤，
空間乾淨、節奏俐落，
卻沒有失去滷肉飯該有的靈魂。
這也是我會推薦給旅客的原因之一：
它保留了臺灣小吃的核心味道，
同時也讓第一次來臺北的人，
吃得安心、坐得舒服。

老臺北胃的帶路小提醒
如果是第一次來：

1. 一定要點招牌黑金滷肉飯
   
2. 可以加一顆滷蛋，風味會更完整
3. 搭配簡單的小菜，就很有臺灣家常感

這不是那種吃完會驚呼「哇！」的料理，
而是會讓你在幾口之後，
慢慢理解
原來，臺灣人的日常，就是這樣被一碗飯照顧著。

地址：103臺北市大同區雙連街55號1樓

電話：0225501379

菜單：https://bio.site/ZhuanTang

第 2 家：富宏牛肉麵｜臺北深夜也醒著的一碗熱湯

如果說滷肉飯代表的是臺灣人的日常，
 那牛肉麵，
 就是很多臺北人心中最有份量的一餐。
而在臺北提到牛肉麵，
 富宏牛肉麵，
 幾乎是夜貓族、加班族、外地旅客一定會被帶去的一站。

為什麼老臺北胃會帶你來吃富宏？
富宏最讓人印象深刻的，
不是華麗裝潢，
而是那鍋永遠冒著熱氣的紅燒湯頭。
湯色濃而不混，
帶著牛骨與醬香慢慢熬出的厚度，
喝起來溫潤、不刺激，
卻會在嘴裡留下很深的記憶點。
牛肉給得大方，
燉到軟嫩卻不鬆散，
搭配彈性十足的麵條，
每一口都很直接、很臺北。

不分時間，任何時候都適合的一碗麵
富宏牛肉麵最迷人的地方，
在於它陪伴了無數個臺北的夜晚。
不管是深夜下班、看完演唱會、
或是剛抵達臺北、還沒適應時差，
這裡總有一碗熱湯在等你。
對旅客來說，
這種不用算時間、不用擔心打烊的安心感，
本身就是一種臺北特色。

老臺北胃的帶路小提醒
第一次來富宏，我會這樣點：

1. 紅燒牛肉麵是首選
   
2. 如果想吃得更過癮，可以加點牛筋或牛肚
3. 湯先喝一口原味，再視情況調整辣度

這不是精緻料理，
卻是一碗能在任何時刻撐住你的牛肉麵。
在臺北，
很多夜晚，
就是靠這樣一碗熱湯走過來的。

地址：108臺北市萬華區洛陽街67號

電話：0223713028

菜單：https://www.facebook.com/pages/富宏牛肉麵-原建宏牛肉麵/

第 3 家：士林夜市・吉彖皮蛋涼麵｜臺北夏天最有記憶點的一口清爽

如果你在夏天來到臺北，
 一定會很快發現一件事
 這座城市，真的很熱。
也正因為這樣，
 臺北的小吃世界裡，
 才會出現像「涼麵」這樣的存在。
而在士林夜市，
 吉彖皮蛋涼麵，
 就是我很常帶旅客來吃的一家。

為什麼在夜市，我會帶你吃涼麵？
很多人對夜市的印象，
都是炸物、熱湯、重口味。
但真正的臺北夜市，
其實也很懂得照顧人的胃。
吉彖的涼麵，
冰涼的麵條拌上濃郁芝麻醬，
再加上切得細緻的皮蛋，
入口的第一瞬間，
就是一種「被降溫」的感覺。
那種清爽，
不是沒味道，
而是在濃香與清涼之間取得剛剛好的平衡。

皮蛋，是靈魂，也是臺灣味的關鍵
對很多外國旅客來說，
皮蛋是既好奇、又有點猶豫的存在。
但我常說，
如果要嘗試皮蛋，
涼麵是一個非常溫柔的起點。
芝麻醬的香氣會先接住味蕾，
皮蛋的風味則在後段慢慢出現，
不衝、不嗆，
反而多了一層深度。
很多人吃完後，
都會露出那種「原來是這樣啊」的表情。

老臺北胃的帶路小提醒
第一次點吉彖皮蛋涼麵，我會建議：

1. 一定要選皮蛋款，才吃得到特色
2. 醬料先拌勻，再吃，風味會更完整
3. 如果天氣真的很熱，這一碗會救你一整晚

這不是華麗的小吃，
卻非常臺北。
在悶熱的夜晚，
站在夜市人潮裡，
吃著一碗涼麵，
你會突然明白——

原來臺北的小吃，連氣候都一起考慮進去了。

地址：111臺北市士林區基河路114號

電話：0981014155

菜單：https://www.facebook.com/profile.php?id=100064238763064

第 4 家：胖老闆誠意肉粥｜臺北人深夜最踏實的一碗粥

如果你問我，
 臺北人在深夜、下班後，
 最容易感到被安慰的食物是什麼——
 我會毫不猶豫地說：肉粥。
而提到肉粥，
 胖老闆誠意肉粥，
 就是很多老臺北人口中的那一味。

為什麼這一碗粥，會被叫做「誠意」？
胖老闆的肉粥，看起來很簡單。
白粥、肉燥、配菜，
沒有華麗擺盤，也沒有複雜作法。
但真正坐下來吃，你會發現：
這碗粥，不敷衍任何一個細節。
粥體滑順、不稀薄，
肉燥香而不膩，
搭配各式家常小菜，
一口一口吃下去，
很自然就會放慢速度。
這種味道，
不是要你驚艷，
而是要你安心。

這不是觀光小吃，而是臺北人的生活片段
胖老闆誠意肉粥，
最迷人的地方，
就是它的客人。
你會看到：

1. 剛下班的上班族
2. 熬夜後來吃一碗熱粥的人
3. 熟門熟路、點菜不用看菜單的老客人

這些畫面，
比任何裝潢都更能說明這家店在臺北的位置。
對旅客來說，
這是一個走進臺北人日常的入口。

老臺北胃的帶路小提醒
第一次來吃，我會這樣建議：

1. 肉粥一定要點，這是主角
2. 配幾樣小菜一起吃，才有完整體驗
3. 不用急，慢慢吃，這碗粥就是要你放鬆

這不是為了拍照而存在的小吃，
而是那種
**會讓人記得「那天晚上，我在臺北吃了一碗很溫暖的粥」**的味道。

地址：10491臺北市中山區長春路89-3號

電話：0913806139

菜單：https://lin.ee/xxbYZyS

第 5 家：圓環邊蚵仔煎｜夜市裡最不能缺席的臺灣經典

如果要選一道
 最常出現在旅客記憶裡的臺灣小吃，
 蚵仔煎一定排得上前幾名。
而在臺北，
 圓環邊蚵仔煎，
 就是那種很多臺北人從小吃到大的存在。

為什麼蚵仔煎，這麼能代表臺灣？
蚵仔煎的魅力，
不在於精緻，
而在於它把幾種看似簡單的食材，
煎成了一種獨特的口感。
新鮮蚵仔的海味、
雞蛋的香氣、
地瓜粉形成的滑嫩外皮，
最後再淋上甜中帶鹹的醬汁，
一口下去，
就是夜市的完整畫面。
這種味道，
很難在其他國家找到替代品。

圓環邊，吃的是記憶感
圓環邊蚵仔煎，
沒有多餘的包裝，
也不刻意迎合潮流。
它留下來的原因很簡單
味道夠穩、節奏夠快、
讓人一吃就知道「對，就是這個」。
對旅客來說，
這是一家
不需要研究、不需要比較，就能安心點蚵仔煎的地方。

老臺北胃的帶路小提醒
第一次吃蚵仔煎，我會這樣建議：

1. 趁熱吃，口感最好
   
2. 不用急著加辣，先吃原味
3. 醬汁是靈魂，別急著把它拌掉

蚵仔煎不是細嚼慢嚥的料理，
它屬於人聲鼎沸、鍋鏟作響的夜市時刻。
站在人群裡，
吃著一盤熱騰騰的蚵仔煎，
你會很清楚地感受到
這，就是臺北的夜晚。

地址：103臺北市大同區寧夏路46號

電話：0225580198

菜單：https://oystera.com.tw/menu

第 6 家：阿淑清蒸肉圓｜第一次吃肉圓，就該從這裡開始

說到臺灣小吃，
 很多人腦中一定會出現「肉圓」兩個字。
但真正吃過之後才會發現，
 肉圓，從來不只有一種樣子。
在臺北，
 阿淑清蒸肉圓，
 就是我很常拿來介紹「清蒸派肉圓」的一家。

清蒸肉圓，和你想像的不一樣
不少旅客對肉圓的第一印象，
來自油炸版本，
外皮厚、口感重。
而阿淑的清蒸肉圓，
完全是另一個方向。
外皮晶瑩、滑嫩，
帶著自然的彈性，
不油、不膩，
一入口反而顯得清爽。
內餡扎實，
豬肉香氣清楚，
搭配特製醬汁，
味道層次簡單卻很乾淨。

為什麼我會推薦給第一次來臺北的旅客？
因為這顆肉圓，
不需要適應期。
它不刺激、不厚重，
即使是第一次嘗試臺灣小吃的人，
也能輕鬆接受。
對旅客來說，
這是一顆
「吃得懂、也記得住」的肉圓。

老臺北胃的帶路小提醒
第一次來阿淑，我會這樣吃：

1. 直接點一顆清蒸肉圓，吃原味
2. 醬汁先別全部拌開，邊吃邊調整
3. 放慢速度，感受外皮的口感變化

這不是夜市裡熱鬧喧囂的料理，
而是那種
安靜地展現臺灣小吃功夫的味道。
當你吃完這顆肉圓，
會更明白一件事
臺灣小吃的魅力，
往往藏在這些細節裡。

地址：242新北市新莊區復興路一段141號

電話：0229975505

第 7 家：胡記米粉湯｜一碗最貼近臺北早晨的味道

如果說前面幾樣小吃，
 是臺北的熱鬧與記憶，
 那麼米粉湯，
 就是這座城市最真實的日常。
而在臺北，
 胡記米粉湯，
 是很多人從小吃到大的存在。

為什麼米粉湯，這麼「臺北」？
米粉湯不是重口味料理，
它靠的不是刺激，
而是一碗清澈卻有深度的湯。
胡記的湯頭，
用豬骨慢慢熬出香氣，
喝起來清爽、不油，
卻能在喉嚨留下溫度。
米粉細軟，
吸附湯汁後入口順滑，
簡單到不能再簡單，
卻正是臺北人習以為常的早晨風景。

配菜，才是這一碗的靈魂延伸
在胡記吃米粉湯，
主角雖然是湯，
但真正讓人滿足的，
往往是那些小菜。
紅燒肉、豬內臟、燙青菜，
隨意點上幾樣，
湯一口、菜一口，
就是很多臺北人記憶中的早餐組合。
對旅客來說，
這是一種
不需要解釋，就能融入的臺北生活感。

老臺北胃的帶路小提醒
第一次來胡記，我會這樣建議：

1. 一定要點米粉湯，湯先喝
2. 再配 1～2 樣小菜，體驗會完整很多
3. 這一餐適合慢慢吃，不用趕

這不是為了觀光而存在的小吃，
而是一碗
每天準時出現在臺北人生活裡的湯。
當你坐在店裡，
聽著湯勺碰撞的聲音，
你會突然感覺到——
原來，臺北的早晨，
就是從這樣一碗米粉湯開始的。

地址：106臺北市大安區大安路一段9號1樓

電話：0227212120

第 8 家：藍家割包｜一口咬下的臺灣街頭記憶

如果要選一道
 外國旅客一看到就會好奇、吃完又會記住的小吃，
 割包，一定在名單裡。
而在臺北，
 藍家割包，
 就是我很放心帶旅客來認識這道經典的一站。

割包，為什麼被叫做「臺灣漢堡」？
割包的結構其實很簡單：
鬆軟的白饅頭、
燉得入味的滷五花肉、
酸菜、花生粉、香菜。
但真正迷人的，
是這些元素組合在一起時，
形成的層次感。
肉香、甜味、鹹味、清爽度，
在一口之間同時出現，
沒有誰搶戲，
卻彼此剛好。
這種平衡感，
正是臺灣小吃很迷人的地方。

藍家割包不是走浮誇路線，
它給人的感覺很直接
就是你期待中的割包樣子。
饅頭柔軟不乾，
五花肉肥瘦比例恰到好處，
入口即化卻不膩口，
花生粉的甜香收尾，
讓整體味道非常完整。
對第一次吃割包的旅客來說，
這是一個
不會出錯、也很容易愛上的版本。

老臺北胃的帶路小提醒
第一次吃藍家割包，我會這樣建議：

1. 直接點招牌割包，不要改配料
2. 如果有香菜，建議保留，味道會更完整
3. 趁熱吃，饅頭口感最好

割包不是精緻料理，
卻非常有記憶點。
站在街頭，
拿著一顆熱騰騰的割包，
邊走邊吃，
你會很清楚地感受到
這一口，就是臺灣的街頭生活。

地址：100臺北市中正區羅斯福路三段316巷8弄3號

電話：0223682060

菜單：https://instagram.com/lan_jia_gua_bao?utm_medium=copy_link

第 9 家：御品元冰火湯圓｜臺北夜晚最溫柔的一碗甜

吃了一整天的臺北小吃，
 到了這個時候，
 胃其實已經差不多滿了。
但只要天氣一涼，
 或夜色慢慢降下來，
 你還是會想找一碗——
 不是為了吃飽，而是為了舒服的甜點。
這時候，我通常會帶你來 御品元冰火湯圓。

為什麼叫「冰火」？這碗湯圓的關鍵就在這裡
御品元最有特色的地方，
就在於它的「冰火交錯」。
熱騰騰的湯圓，
外皮軟糯、內餡濃香，
搭配冰涼清甜的桂花蜜湯，
一口下去，
溫度在嘴裡交替出現。
不是衝突，
而是一種很細膩的平衡。
這樣的吃法，
也正是臺灣甜點很擅長的地方——
不張揚，但很有記憶點。

這是一碗，會讓人慢下來的甜點
和夜市裡熱鬧的甜品不同，
御品元的冰火湯圓，
更像是一個讓人停下腳步的存在。
你會發現，
坐在這裡吃湯圓的人，
說話聲都會不自覺地變小。
對旅客來說，
這不只是吃甜點，
而是一個
把白天的熱鬧慢慢收進回憶裡的時刻。

老臺北胃的帶路小提醒
第一次吃御品元，我會這樣建議：

1. 點招牌冰火湯圓，體驗完整特色
2. 先單吃湯圓，再搭配湯一起吃
3. 放慢速度，這一碗不適合趕時間

這不是為了拍照而存在的甜點，
而是一碗
會讓你記得「那天晚上在臺北，很舒服」的湯圓。

地址：106臺北市大安區通化街39巷50弄31號

電話：0955861816

菜單：https://instagram.com/lan_jia_gua_bao

第 10 家：頃刻間綠豆沙牛奶專賣店｜把臺北的味道，留在最後一口清甜

走到這一站，
 其實已經不需要再吃什麼大份量的東西了。
這時候，
 最適合的，
 是一杯不吵鬧、不張揚，
 卻會默默留在記憶裡的飲品。
頃刻間綠豆沙牛奶，
 就是我很常用來替一天畫下句點的選擇。

綠豆沙牛奶，為什麼這麼「臺灣」？
在臺灣，
飲料不只是解渴，
而是一種生活節奏。
綠豆沙牛奶看起來簡單，
但真正好喝的版本，
靠的是火候、比例，
還有耐心。
頃刻間的綠豆沙，
口感細緻、不粗顆，
甜度自然、不膩口，
牛奶的加入，
讓整杯變得柔順而溫和。
這不是衝擊味蕾的飲料，
而是一種
喝完之後，會覺得剛剛那一刻很舒服的甜。

為什麼我會用它當作最後一站？
因為它很臺北。
你可以外帶，
邊走邊喝；
也可以站在店門口，
慢慢把杯子喝空。
沒有儀式感，
卻很真實。
對旅客來說，
這杯綠豆沙牛奶，
就像是把今天吃過的所有味道，
溫柔地整理好，
帶走。

老臺北胃的帶路小提醒
第一次喝頃刻間，我會這樣建議：

1. 直接點招牌綠豆沙牛奶
   
2. 正常甜就很剛好，不用特別調整
3. 找個角落慢慢喝，別急著趕路

這一杯，
不會讓你驚呼，
卻會在回程的路上，
突然想起來。
原來，臺北的味道，是這樣結束一天的。

地址：111臺北市士林區小北街1號

電話：0228818619

菜單：https://instagram.com/chill_out_moment?igshid=YmMyMTA2M2Y=

如果只有 3 天的自助旅行在臺北，怎麼吃這 10 家？

第一次來臺北，
時間有限、胃容量也有限，
與其每一家都趕，不如照著節奏吃。
這份 3 天小吃路線，
是老臺北胃會帶朋友實際走的版本：
不爆走、不硬塞，
讓你每天都吃得剛剛好。

臺北 3 天小吃推薦行程表（老臺北胃版本）

天數	時段	店家名稱	小吃類型
Day 1	午餐	饌堂－黑金滷肉飯（雙連店）	滷肉飯
Day 1	下午	阿淑清蒸肉圓	肉圓
Day 1	晚餐	富宏牛肉麵	牛肉麵
Day 1	宵夜	胖老闆誠意肉粥	粥品
Day 2	早餐	胡記米粉湯	米粉湯
Day 2	下午	藍家割包	割包
Day 2	晚上	士林夜市－吉彖皮蛋涼麵	涼麵
Day 2	夜市	圓環邊蚵仔煎	蚵仔煎
Day 3	下午	御品元冰火湯圓	甜點
Day 3	收尾	頃刻間綠豆沙牛奶專賣店	飲品

雖然每個小吃的地點都有一點距離，但是你也知道，好吃的小吃，是值得你花一點時間前往品嘗
老臺北胃的小提醒

1. 不需要每一家都點到最滿
2. 留一點餘裕，才會想再回來
3. 臺北小吃的魅力，不在於吃多少，而在於記住了什麼味道
   

當你照著這 3 天走完，
你會發現，
臺北不是靠一兩道名菜被記住的，
而是靠這些看似日常、卻很真實的小吃。
下次再來，老臺北胃再帶你吃更深的那一輪。

老臺北胃帶路｜這 10 口，就是我心中的臺北

寫到這裡，
 其實已經不是在推薦哪一家小吃了。
而是在回頭看，
 這座城市，是怎麼用食物陪著人生活的。
滷肉飯、牛肉麵、肉粥、米粉湯，
 不是為了成為觀光名單而存在，
 而是每天默默出現在臺北人的日子裡。
夜市裡的蚵仔煎、涼麵、割包，
 熱鬧、吵雜、節奏很快，
 卻也正是臺北最真實的樣子。
而最後那碗湯圓、那杯綠豆沙牛奶，
 則是在一天結束時，
 替味蕾留下一個溫柔的句點。

如果你問我，
「這 10 家是不是臺北最好吃的小吃？」
我會說，
它們不一定是排行榜第一名，
卻是我真的會帶朋友去吃的版本。
因為它們吃得到：

1. 臺北人的日常
2. 巷弄裡的熟悉感
3. 不需要解釋，就能被理解的味道

如果你是第一次來臺北，
跟著這份清單走，
你不一定會吃得最飽，
但你一定會記得——
臺北，是什麼味道。
而如果有一天，
你又再回到這座城市，
走進熟悉的街口、
看到冒著熱氣的小攤，
你也會開始懂得，
為什麼老臺北胃，
總是記得這些看似平凡的滋味。
因為，真正留在心裡的，
從來不是吃過多少，
而是哪一口，讓你想起臺北。

御品元冰火湯圓招牌值得嗎？

走完這 10 家，

你可能會發現一件事胖老闆誠意肉粥值得一試嗎？

臺北的小吃，其實不急著被你記住。

它們就安靜地存在在街角、夜市、轉彎處，阿淑清蒸肉圓CP 值高嗎？

等你有一天，再回到這座城市。富宏牛肉麵招牌值得嗎？

如果你是第一次來臺北，富宏牛肉麵適合第一次吃嗎？

希望這份「老臺北胃帶路」的清單，

能幫你少一點猶豫、多一點安心。

不用擔心踩雷，胖老闆誠意肉粥不加辣好吃嗎？

也不用為了排行而奔波，阿淑清蒸肉圓女生會喜歡嗎？

只要照著節奏走，

你就會吃到屬於自己的臺北味道。

而如果你已經來過臺北，

那更希望這篇文章，頃刻間綠豆沙牛奶專賣店份量有誠意嗎？

能帶你走進那些

你可能錯過、卻一直都在的日常小吃。

因為真正迷人的旅行，

從來不是把清單全部打勾，

而是某一天，

你突然想起那碗飯、那口湯、那杯甜，饌堂－黑金滷肉飯（雙連店）會不會太鹹？

然後在心裡對自己說一句：胖老闆誠意肉粥值得排隊嗎？

「下次再去臺北，還想再吃一次。」

把這篇文章存起來、分享給一起旅行的人，

或是在規劃行程時，再回來看看。

讓味道，成為你認識臺北的方式。

下一次來臺北，

別急著走遠。

老臺北胃，饌堂－黑金滷肉飯（雙連店）冬天適合吃嗎？

會一直在這些地方，

等你再回來。

Diatoms (blue/white/yellow) frozen on an electron microscopy grid (copper) during a sample preparation step for cryo-electron tomography. Credit: Benoit Gallet and Martin Oeggerli, Micronaut A groundbreaking study reveals a protein shell in diatoms that enhances their CO2 fixation capabilities, offering new avenues for bioengineering to combat climate change by optimizing photosynthesis. Tiny ocean diatoms are highly efficient at capturing carbon dioxide (CO2) from the environment, accounting for up to 20 percent of the Earth’s CO2 fixation. Researchers at the University of Basel in Switzerland have now discovered a protein shell within these algae that is essential for their ability to fix CO2 so effectively. This significant finding could inspire new bioengineering strategies to help reduce atmospheric CO2 levels. Diatom Discovery and Carbon Capture Diatoms, though invisible to the naked eye, are among the most productive algae in the ocean and play a crucial role in the global carbon cycle. Through photosynthesis, they absorb large amounts of CO2 from the environment and convert it into nutrients that sustain much of ocean life. Despite their significance, how diatoms perform this process so efficiently has remained a mystery. Now, researchers led by Prof. Ben Engel at the University of Basel’s Biozentrum, along with teams from the University of York, UK, and Kwansei-Gakuin University in Japan, have uncovered a protein shell crucial to diatoms’ CO2 fixation. Using advanced imaging techniques like cryo-electron tomography (cryo-ET), they mapped the molecular structure of the PyShell protein sheath and revealed its function. These findings were recently published in two papers in the journal Cell. Cryo-electron tomography reveals the molecular architecture of the diatom pyrenoid, showing how Rubisco is surrounded by the PyShell, increasing the efficiency of carbon dioxide fixation. Credit: Gröger et al. / Manon Demulder, Biozentrum, Universität Basel PyShell: A Key to Efficient Photosynthesis In plants and algae, photosynthesis takes place in chloroplasts. Inside these chloroplasts, energy from sunlight is harvested by thylakoid membranes and then used to help the enzyme Rubisco fix CO2. However, algae have an advantage: they pack all their Rubisco into small compartments called pyrenoids, where CO2 can be captured more efficiently. “We have now discovered that diatom pyrenoids are encased in a lattice-like protein shell,” says Dr. Manon Demulder, author on both studies. “The PyShell not only gives the pyrenoid its shape, but it helps create a high CO2 concentration in this compartment. This enables Rubisco to efficiently fix CO2 from the ocean and convert it into nutrients.” When the researchers removed the PyShell from the algae, their ability to fix CO2 was significantly impaired. Photosynthesis and cell growth were reduced. “This showed us how important the PyShell is for efficient carbon capture – a process that is crucial for ocean life and the global climate,” says Manon Demulder. Potential Climate Solutions Through Bioengineering The discovery of the PyShell could also open promising avenues for biotechnological research aimed at combatting climate change – one of the most pressing challenges of our time. “First of all, we humans must reduce our CO2 emissions to slow the pace of climate change. This requires immediate action,” says Ben Engel. “The CO2 that we emit now will remain in our atmosphere for thousands of years. We hope that discoveries such as the PyShell can help inspire new biotechnology applications that improve photosynthesis and capture more CO2 from the atmosphere. These are long-term goals, but given the irreversibility of CO2 emissions, it is important that we perform basic research now to create more opportunities for future carbon-capture innovations.” Reference: “Diatom pyrenoids are encased in a protein shell that enables efficient CO2 fixation” by Ginga Shimakawa, Manon Demulder, Serena Flori, Akihiro Kawamoto, Yoshinori Tsuji, Hermanus Nawaly, Atsuko Tanaka, Rei Tohda, Tadayoshi Ota, Hiroaki Matsui, Natsumi Morishima, Ryosuke Okubo, Wojciech Wietrzynski, Lorenz Lamm, Ricardo D. Righetto, Clarisse Uwizeye, Benoit Gallet, Pierre-Henri Jouneau, Christoph Gerle, Genji Kurisu, Giovanni Finazzi, Benjamin D. Engel and Yusuke Matsuda, 1 October 2024, Cell. DOI: 10.1016/j.cell.2024.09.013 “A protein blueprint of the diatom CO2-fixing organelle” by Onyou Nam, Sabina Musiał, Manon Demulder, Caroline McKenzie, Adam Dowle, Matthew Dowson, James Barrett, James N. Blaza, Benjamin D. Engel and Luke C.M. Mackinder, 4 October 2024, Cell. DOI: 10.1016/j.cell.2024.09.025

Researchers discovered the “Octopus Garden,” a deep-sea nursery off the Central California coast where octopuses mate and nest, benefiting from hydrothermal springs’ warmth. Although protected, further conservation efforts are needed to shield these unique deep-sea habitats from human threats. Credit: © 2022 MBARI MBARI’s advanced technology offers new insight into the “Octopus Garden” off Central California, the largest aggregation of octopus on Earth. In 2018, researchers from NOAA’s Monterey Bay National Marine Sanctuary and Nautilus Live observed thousands of octopus nesting on the deep seafloor off the Central California coast. The discovery of the “Octopus Garden” captured the curiosity of millions of people around the world, including MBARI scientists. For three years, MBARI and collaborators used high-tech tools to monitor the Octopus Garden and learn exactly why this site is so attractive for deep-sea octopus. Purpose of the Garden and Its Unique Properties In a new study published today (August 23) in the journal Science Advances, a team of researchers from MBARI, NOAA’s Monterey Bay National Marine Sanctuary, Moss Landing Marine Laboratories, the University of Alaska Fairbanks, the University of New Hampshire, and the Field Museum confirmed that deep-sea octopus migrate to the Octopus Garden to mate and nest. The Octopus Garden is one of a handful of known deep-sea octopus nurseries. At this nursery, warmth from deep-sea thermal springs accelerates the development of octopus eggs. Scientists believe the shorter brooding period increases a hatchling octopus’ odds for survival. The Octopus Garden is the largest known aggregation of octopus on the planet—researchers counted more than 6,000 octopus in a portion of the site and expect there may be 20,000 or more at this nursery. “Thanks to MBARI’s advanced marine technology and our partnership with other local researchers, we were able to observe the Octopus Garden in tremendous detail, which helped us discover why so many deep-sea octopus gather there. These findings can help us understand and protect other unique deep-sea habitats from climate impacts and other threats,” said MBARI Senior Scientist Jim Barry, lead author of the new study. An aggregation of female pearl octopus (Muusoctopus robustus) nesting at the Octopus Garden, located near Davidson Seamount off the Central California at a depth of approximately 3,200 meters. Researchers used MBARI’s advanced technology to confirm pearl octopus gather at the Octopus Garden to mate and nest. Warm water from hydrothermal springs accelerates development of octopus embryos, giving young octopus a better chance of survival. Credit: © 2022 MBARI Location and Behavior of the Octopuses The Octopus Garden is located 3,200 meters (10,500 feet, or about two miles) below the ocean’s surface on a small hill near the base of Davidson Seamount, an extinct underwater volcano 130 kilometers (80 miles) southwest of Monterey, California. The site is full of Muusoctopus robustus—a species MBARI researchers nicknamed the pearl octopus because from a distance, nesting individuals look like opalescent pearls on the seafloor. Over the course of 14 dives with MBARI’s remotely operated vehicle (ROV) Doc Ricketts, the research team learned why such large numbers of pearl octopus are attracted to this location. The presence of adult male and female octopus, developing eggs, and octopus hatchlings indicated that the site is used exclusively for reproduction. The team did not observe any intermediate-sized individuals or any evidence of feeding. Pearl octopus gather at this site solely to mate and nest. When researchers from NOAA and Nautilus Live first discovered the Octopus Garden, they observed “shimmering” waters. This phenomenon occurs when warm and cool waters mix, suggesting the region had previously unknown thermal springs. Further investigation by MBARI researchers and their collaborators confirmed octopus nests are clustered in crevices bathed by hydrothermal springs where warmer waters flow from the seafloor. Impact of Temperature on Octopus Development The ambient water temperature at 3,200 meters (10,500 feet) deep is 1.6 degrees Celsius (about 35 degrees Fahrenheit). However, the water temperature within the cracks and crevices at the Octopus Garden reaches nearly 11 degrees Celsius (about 51 degrees Fahrenheit). Octopuses are ectotherms, or cold-blooded animals. The frigid temperatures of the deep sea slow their metabolism as well as their rate of embryonic development. Most deep-sea octopuses have very long incubation periods compared to their relatives inhabiting warmer shallow seas. Past experiments have measured egg incubation time for a number of octopus species in habitats and locations around the world. Comparing those egg incubation times clearly demonstrates how temperature affects the rate of embryo development—the colder the water, the slower the embryos grow. At the near-freezing temperatures of the abyss, researchers expected pearl octopus eggs to take five to eight years, if not longer, to hatch. A 4K camera on MBARI’s ROV Doc Ricketts provided a close-up look at nesting mothers. MBARI researchers and their collaborators used the scars and other distinguishing features of individual octopus moms to monitor the development of their broods. Surprisingly, the eggs hatched in less than two years. Warmth from thermal springs increased the metabolism of female octopus and their broods, reducing the time required for incubation. Researchers believe the shorter brood period in warmer waters greatly reduces the risk that developing octopus embryos will be injured or eaten by predators. Nesting in warmer water boosts the reproductive success of the pearl octopus, better ensuring the offspring’s survival. “The deep sea is one of the most challenging environments on Earth, yet animals have evolved clever ways to cope with frigid temperatures, perpetual darkness, and extreme pressure. Very long brooding periods increase the likelihood that a mother’s eggs won’t survive. By nesting at hydrothermal springs, octopus moms give their offspring a leg up,” said Barry. Ecology and Significance of the Garden The massive number of octopus in one area attracts both predators and scavengers. Like most other cephalopods, pearl octopus die after they reproduce. Dead octopus at the Octopus Garden provide a feast for scavengers. A rich community of invertebrates lives alongside the nesting females, undoubtedly benefiting from unhatched eggs, vulnerable hatchlings, or adult octopus that have died. Davidson Seamount and its Octopus Garden are protected as part of Monterey Bay National Marine Sanctuary. Previous MBARI expeditions to Davidson Seamount in 2002 and 2006 revealed the stunning community of life on its rocky slopes. MBARI’s images and video of beautiful deep-sea corals, vibrant sponges, and curious fishes engaged and inspired audiences worldwide. Ocean champions spoke up to protect this unique, and still untouched, ocean wilderness. In 2008, resource managers expanded the Monterey Bay National Marine Sanctuary to include Davidson Seamount. “Essential biological hotspots like this deep-sea nursery need to be protected,” said Barry. “Climate change, fishing, and mining threaten the deep sea. Protecting the unique environments where deep-sea animals gather to feed or reproduce is critical, and MBARI’s research is providing the information that resource managers need for decision-making.” This work is funded as part of the David and Lucile Packard Foundation’s long-term support of MBARI’s ocean research and technology. Deep-Sea Exploration and Monitoring For more than two decades, researchers from MBARI and NOAA have collaborated to study Davidson Seamount. Since the first expedition to the seamount in 2002, NOAA has leveraged MBARI expertise in marine geology and benthic biology and ecology to develop a comprehensive research program that aims to understand the unique community of life on and around Davidson Seamount. Now, Davidson Seamount is considered one of the best-studied and well-protected seamounts in the world. In October 2018, a team of researchers from NOAA, the Ocean Exploration Trust, and collaborators made an expedition to Davidson Seamount aboard the E/V Nautilus. At the suggestion of MBARI geologists and NOAA researchers, the Nautilus Live team decided to expand their exploration from the top of the seamount to its surrounding foothills. The researchers discovered thousands of octopus aggregated around a rocky ridge adjacent to the towering seamount. Most of the octopus were oriented upside down, inverting their arms and folding them around their bodies. This posture was an indication of pearl octopus (Muusoctopus robustus) mothers protecting, or brooding, their eggs. The pearl octopus is a pale purple species about the size of a grapefruit that occurs in the northeastern Pacific Ocean from Oregon to Baja California. MBARI has observed this species at depths of 2,300 to 3,600 meters (7,500 to 11,800 feet). MBARI researchers and their collaborators deployed a suite of advanced scientific instruments developed by MBARI engineers to better understand the Octopus Garden. “The expertise of the MBARI team—the engineers, pilots of our submersible vehicles, and crew of our research vessels—was integral to studying this hotspot of life two miles below the surface. We leveraged decades of experience in deep-sea exploration to develop and deploy instruments to study the Octopus Garden without disturbing the nesting mothers,” said Barry. MBARI’s ROV Doc Ricketts recorded high-definition and 4K video of the brooding pearl octopus and their neighbors. MBARI’s skilled submersible pilots maneuvered the ROV close to brooding pearl octopus to deploy instruments to measure the environmental conditions within their nests, including temperature and oxygen levels, and to film mothers and their eggs up close in ultra-high definition resolution. A stereoscopic camera allowed MBARI engineers to visualize sites within the Octopus Garden in 3D. The team also launched one of MBARI’s autonomous underwater vehicles to map the Octopus Garden at meter-scale resolution. MBARI engineers outfitted the ROV Doc Ricketts with an innovative, custom-built sensor suite, the Low-Altitude Survey System (LASS), to see the Octopus Garden in even greater detail. The LASS gathered detailed bathymetry information to help researchers characterize the seafloor habitat at centimeter-scale resolution. The LASS also took high-resolution photographs of the Octopus Garden. Researchers assembled these photographs into a photomosaic to count the number of nests within this deep-sea nursery. They documented 5,718 octopus within a 2.5-hectare (6.2-acre) area at the center of the Octopus Garden. The team estimated the total population of the 333-hectare (823-acre) hillock could easily exceed 20,000 individuals. A time-lapse camera collected long-term observations of the octopus’ behavior and changes in the community over a period of more than six months, allowing researchers to keep watch on the octopus nursery between research expeditions. The camera recorded an image every 20 minutes and amassed a trove of more than 12,200 images from March 2022 to August 2022. These photographs revealed various activities and behaviors of octopus, their predators, and local scavengers. Both male and female pearl octopus migrate to the Octopus Garden. Females search for a warm nesting spot to deposit a clutch of approximately 60 elongate, sausage-shaped eggs. When brooding, mothers cover their eggs with their body and protect them from predators that creep too close. She lives off food reserves from her own tissues while tending to her developing eggs. The transformation from egg to hatchling is not easy. In addition to going through development successfully, embryos must avoid injury, predation, infection, and other external sources of mortality. Maternal care protects them from most external risks, but a shorter brooding period generally allows more eggs to survive. As is typical of cephalopods, male and female pearl octopus die after reproducing—the Octopus Garden will be their final resting spot. Most females live until their eggs have hatched. Sometimes, however, a mother octopus runs out of energy and dies before her eggs complete their development, exposing the developing eggs to greater risk. The time-lapse camera revealed that nesting mothers push aside the carcasses of dead octopus. Food is scarce in the deep sea and nothing goes to waste. Larger scavengers like rattail fishes (family Macrouridae), cusk eels (family Ophidiidae), whelks, and sea anemones feast on octopus remains. Near Davidson Seamount, life on the deep seafloor depends on the rain of organic matter from above. Researchers estimated the turnover of male octopus and nesting females to calculate how much nutrition this massive aggregation provides. Biomass from dying octopus represents a substantial carbon subsidy to the local seafloor community, providing 72 percent more food than is available outside the Octopus Garden. Challenges and Need for Protection Many questions still remain about the Octopus Garden, including where pearl octopus go after hatching, how this octopus species became adapted to breeding in thermal springs, how adult octopus find the thermal springs, what advantage individuals breeding in these hydrothermal springs have over those that breed elsewhere, and how common hydrothermal springs are in the deep sea. The deep sea is not immune to threats like fishing, pollution, and climate change. By documenting deep-sea biodiversity and identifying hotspots of life on the ocean floor, scientists are gathering important information that resource managers can use to guide protections for this unique environment and its inhabitants. “Technological advances in our ability to study the ocean have helped us discover and document incredible biodiversity across an array of deep-sea environments. As the imprint of human activities reaches deeper into ocean ecosystems, we need to protect not only the octopus nurseries found off California and Costa Rica, but also the many other biological treasures that remain undiscovered,” emphasized Barry. Deep-sea octopus nurseries: A new field of exploration Researchers have documented four deep-sea octopus nurseries to date—two off the coast of Central California and two off the coast of Costa Rica—and are continuing to study these sites to learn more about octopus behavior. December 2013: Discovery of first octopus nursery at Dorado Outcrop (Costa Rica) Researchers from the University of Akron, the Field Museum, and the University of Alaska Fairbanks observed an aggregation of more than 100 octopus at the Dorado Outcrop, a hydrothermal spring located approximately 160 kilometers (100 miles) off the Pacific coast of Costa Rica at a depth of 3,000 meters (9,800 feet). The team identified the octopus as a potentially undescribed species of Muusoctopus. Nearly all of the individuals were in a brooding position, however, none of the eggs that researchers observed were viable. April 2018: Researchers publish findings from the Dorado Outcrop (Costa Rica) The team of researchers from the University of Akron, the Field Museum, and the University of Alaska Fairbanks published their observations of deep-sea octopus brooding unviable eggs at the Dorado Outcrop in Deep Sea Research Part I. October 2018: Discovery of second octopus nursery at the Octopus Garden (Davidson Seamount, United States) During a Nautilus Live expedition with the E/V Nautilus, researchers from NOAA’s Monterey Bay National Marine Sanctuary, the Ocean Exploration Trust, and collaborators observed a large aggregation of brooding octopus on a hillock approximately 12 kilometers (7.5 miles) southeast of Davidson Seamount at a depth of 3,200 meters (10,500 feet). Researchers identified the octopus as Muusoctopus robustus. A second visit by researchers from NOAA and the Woods Hole Oceanographic Institution (WHOI) in March 2019 confirmed the presence of warm hydrothermal springs at this site. The expedition team also confirmed that the octopus were brooding viable eggs and observed baby octopus hatching from the eggs. April 2019: First MBARI expedition to Octopus Garden (Davidson Seamount, United States) MBARI researchers made their first visit to the Octopus Garden as part of the 2019 Seafloor Ecology expedition. Along with collaborators, they visited the site 14 times with the R/V Western Flyer between April 2019 and August 2022. Additionally, MBARI researchers visited the Octopus Garden with the R/V Rachel Carson in February 2022 to launch a mapping autonomous underwater vehicle and create meter-scale maps of the site. October 2019: Discovery of third octopus nursery at Octocone (Davidson Seamount, United States) During a Nautilus Live expedition with the E/V Nautilus, researchers from NOAA, the Ocean Exploration Trust, and collaborators observed a second aggregation of brooding octopus on a volcanic cone to the east of Davidson Seamount. This site is approximately 17 kilometers (10.5 miles) northeast of the Octopus Garden. Researchers identified the octopus as Muusoctopus robustus. The octopus were confirmed to be brooding viable eggs. June 2023: Discovery of fourth octopus nursery (Costa Rica) During a Schmidt Ocean Institute expedition with the R/V Falkor (too), researchers from the Bigelow Laboratory for Ocean Sciences and the University of Costa Rica observed a previously unknown octopus nursery near an unexplored and still-unnamed seamount off the Pacific coast of Costa Rica. Upon returning to the nearby Dorado Outcrop, the team also observed octopus brooding viable eggs, confirming this location is indeed an active octopus nursery. Both Costa Rican nurseries host a potentially undescribed species of Muusoctopus. August 2023: MBARI researchers publish findings from the Octopus Garden (Davidson Seamount, United States) MBARI researchers and their collaborators from NOAA, Moss Landing Marine Laboratories, the University of Alaska Fairbanks, the University of New Hampshire, and the Field Museum published their research on brooding pearl octopus in Science Advances, confirming that deep-sea octopus migrate to the Octopus Garden to mate and nest. Reference: “Abyssal hydrothermal springs—Cryptic incubators for brooding octopus” by James P. Barry, Steven Y. Litvin, Andrew DeVogelaere, David W. Caress, Chris F. Lovera, Amanda S. Kahn, Erica J. Burton, Chad King, Jennifer B. Paduan, C. Geoffrey Wheat, Fanny Girard, Sebastian Sudek, Anne M. Hartwell, Alana D. Sherman, Paul R. McGill, Aaron Schnittger, Janet R. Voight and Eric J. Martin, 23 August 2023, Science Advances. DOI: 10.1126/sciadv.adg3247

Scripps Research scientists have developed a simpler method to add new amino acids to proteins using four-nucleotide codons, creating novel peptides with potential applications in drug discovery and beyond. Credit: SciTechDaily.com Scripps Research scientists have created a method using four-nucleotide codons to incorporate non-canonical amino acids into proteins, expanding protein engineering possibilities without requiring genome-wide edits. This method has been tested in creating new peptides and holds potential for applications in various fields. In every introductory biology class, it’s a fundamental concept: proteins are made from combinations of 20 distinct amino acids, arranged in various sequences like words. However, researchers aiming to engineer biological molecules with novel functions have long found these 20 building blocks restrictive. As a result, they have sought ways to incorporate new components—known as non-canonical amino acids—into proteins. Now, scientists at Scripps Research have designed a new paradigm for easily adding non-canonical amino acids to proteins. Their approach, described in Nature Biotechnology on September 11, 2024, revolves around using four RNA nucleotides—rather than the typical three—to encode each new amino acid. A New Approach to Protein Engineering “Our goal is to develop proteins with tailored functions for applications in fields spanning bioengineering to drug discovery,” says senior author Ahmed Badran, PhD, an assistant professor of chemistry at Scripps Research. “Being able to incorporate non-canonical amino acids into proteins with this new method gets us closer to that goal.” For a cell to produce any given protein, it must translate a strand of RNA into a string of amino acids. Every three nucleotides of RNA, called a codon, correspond to one amino acid. But many amino acids have more than one possible codon; for instance, RNA reading the sequences UAU and UAC both correspond to the amino acid tyrosine. It’s the job of small molecules called transfer RNAs (tRNAs) to link each amino acid to its corresponding codons. Examples of the >100 macrocycles generated in this study. Colored components represent new-to-nature amino acids that were incorporated into either peptide. Credit: Scripps Research Recently, researchers aiming to add completely new amino acids to a protein have created strategies to reassign a codon. For instance, the UAU codon could be linked to a new amino acid by changing the tRNA for UAU; this would result in UAU being read by the cell as corresponding to a building block other than tyrosine. But at the same time, every instance of UAU in the cell’s genome would need to become UAC, in order to prevent the new amino acid from being integrated into thousands of other proteins where it doesn’t belong. “Creating free codons by whole genome recoding can be a powerful strategy, but it can also be a challenging undertaking since it requires considerable resources to build new genomes,” says Badran. “For the organism itself, it can be difficult to predict how such codon changes influence genome stability and host protein production.” Introducing Four-Nucleotide Codons Badran and his colleagues wanted to create an efficient plug-and-play strategy that would only incorporate the chosen non-canonical amino acid(s) into specific sites in a target protein, without disrupting the cell’s normal biology or requiring the entire genome to be edited. That meant using tRNA that wasn’t already assigned to an amino acid. Their solution: a four-nucleotide codon. The team knew that in a few situations—such as bacteria quickly adapting to resist drugs—four-nucleotide codons had naturally evolved. So, in their new work, the researchers studied what caused cells to use a codon with four nucleotides rather than three. They discovered that the identities of the sequences nearby to the four-base codon were critical—frequently used codons enhanced how the cell could read a four-nucleotide codon to incorporate a non-canonical amino acid. Badran’s group then tested whether they could alter the sequence of a single gene so that it had a new four-nucleotide codon that would be correctly used by the cell. The method worked: When the researchers surrounded a target site with three-letter, frequently used codons and maintained sufficient levels of the four-nucleotide tRNA, the cell incorporated any new amino acid that was attached to the corresponding four-letter tRNA. The research team repeated the experiment with 12 different four-nucleotide codons and then used the technique to design more than 100 new cyclic peptides—called macrocycles—with up to three non-canonical amino acids in each. “These cyclic peptides are reminiscent of bioactive small molecules that one might find in nature,” says Badran. “By capitalizing on the programmability of protein synthesis and the diversity of building blocks accessible by this approach, we can create new-to-nature small molecules that will have exciting applications in drug discovery.” Benefits of the New Method He adds that, compared with previous approaches to non-canonical amino acid incorporation, this new method is easy to use since it involves altering only one gene rather than a cell’s entire genome. Additionally, more non-canonical amino acids could be used in a single protein since there are more possible four-nucleotide codons than three-nucleotide ones. “Our results suggest that one can now easily and effectively incorporate non-canonical amino acids at diverse sites in a wide array of proteins,” says Badran. “We’re excited about these possibilities for our ongoing work and to provide this capability to the broader community.” He notes that the technique could be used to re-engineer existing proteins—or create entirely new ones—that have utility in a range of sectors, including medicine, manufacturing and chemical sensing. Reference: “Efficient genetic code expansion without host genome modifications” by Alan Costello, Alexander A. Peterson, David L. Lanster, Zhiyi Li, Gavriela D. Carver and Ahmed H. Badran, 11 September 2024, Nature Biotechnology. DOI: 10.1038/s41587-024-02385-y This work was supported by funding from the National Institutes of Health (DP5-OD024590), the Research Corporation for Science Advancement, the Sloan Foundation (G-2023-19625), the Thomas Daniel Innovation Fund (627163_1), an Abdul Latif Jameel Water and Food Systems Lab Grand Challenge Award (GR000141-S6241), a Breakthrough Energy Explorer Grant (GR000056), the Foundation for Food & Agriculture Research (28-000578), a Homeworld Collective Garden Grant (GR000129), the Army Research Office (81341- BB-ECP), the Hope Funds for Cancer Research (HFCR-23-03-01), a Skaggs-Oxford Scholarship and a Fletcher Jones Foundation Fellowship.

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